CINXE.COM

<!DOCTYPE html> <html lang="en" class="no-js"> <head> <meta charset="UTF-8"> <meta http-equiv="X-UA-Compatible" content="IE=edge"> <meta name="applicable-device" content="pc,mobile"> <meta name="viewport" content="width=device-width, initial-scale=1"> <meta name="robots" content="max-image-preview:large"> <meta name="access" content="Yes"> <meta name="360-site-verification" content="1268d79b5e96aecf3ff2a7dac04ad990" /> <title>Fish do not feel pain and its implications for understanding phenomenal consciousness | Biology & Philosophy</title> <meta name="twitter:site" content="@SpringerLink"/> <meta name="twitter:card" content="summary_large_image"/> <meta name="twitter:image:alt" content="Content cover image"/> <meta name="twitter:title" content="Fish do not feel pain and its implications for understanding phenomenal consciousness"/> <meta name="twitter:description" content="Biology & Philosophy - Phenomenal consciousness or the subjective experience of feeling sensory stimuli is fundamental to human existence. Because of the ubiquity of their subjective..."/> <meta name="twitter:image" content="https://media.springernature.com/full/springer-static/cover-hires/journal/10539"/> <meta name="journal_id" content="10539"/> <meta name="dc.title" content="Fish do not feel pain and its implications for understanding phenomenal consciousness"/> <meta name="dc.source" content="Biology & Philosophy 2014 30:2"/> <meta name="dc.format" content="text/html"/> <meta name="dc.publisher" content="Springer"/> <meta name="dc.date" content="2014-12-16"/> <meta name="dc.type" content="OriginalPaper"/> <meta name="dc.language" content="En"/> <meta name="dc.copyright" content="2014 The Author(s)"/> <meta name="dc.rights" content="2014 The Author(s)"/> <meta name="dc.rightsAgent" content="journalpermissions@springernature.com"/> <meta name="dc.description" content="Phenomenal consciousness or the subjective experience of feeling sensory stimuli is fundamental to human existence. Because of the ubiquity of their subjective experiences, humans seem to readily accept the anthropomorphic extension of these mental states to other animals. Humans will typically extrapolate feelings of pain to animals if they respond physiologically and behaviourally to noxious stimuli. The alternative view that fish instead respond to noxious stimuli reflexly and with a limited behavioural repertoire is defended within the context of our current understanding of the neuroanatomy and neurophysiology of mental states. Consequently, a set of fundamental properties of neural tissue necessary for feeling pain or experiencing affective states in vertebrates is proposed. While mammals and birds possess the prerequisite neural architecture for phenomenal consciousness, it is concluded that fish lack these essential characteristics and hence do not feel pain."/> <meta name="prism.issn" content="1572-8404"/> <meta name="prism.publicationName" content="Biology & Philosophy"/> <meta name="prism.publicationDate" content="2014-12-16"/> <meta name="prism.volume" content="30"/> <meta name="prism.number" content="2"/> <meta name="prism.section" content="OriginalPaper"/> <meta name="prism.startingPage" content="149"/> <meta name="prism.endingPage" content="165"/> <meta name="prism.copyright" content="2014 The Author(s)"/> <meta name="prism.rightsAgent" content="journalpermissions@springernature.com"/> <meta name="prism.url" content="https://link.springer.com/article/10.1007/s10539-014-9469-4"/> <meta name="prism.doi" content="doi:10.1007/s10539-014-9469-4"/> <meta name="citation_pdf_url" content="https://link.springer.com/content/pdf/10.1007/s10539-014-9469-4.pdf"/> <meta name="citation_fulltext_html_url" content="https://link.springer.com/article/10.1007/s10539-014-9469-4"/> <meta name="citation_journal_title" content="Biology & Philosophy"/> <meta name="citation_journal_abbrev" content="Biol Philos"/> <meta name="citation_publisher" content="Springer Netherlands"/> <meta name="citation_issn" content="1572-8404"/> <meta name="citation_title" content="Fish do not feel pain and its implications for understanding phenomenal consciousness"/> <meta name="citation_volume" content="30"/> <meta name="citation_issue" content="2"/> <meta name="citation_publication_date" content="2015/03"/> <meta name="citation_online_date" content="2014/12/16"/> <meta name="citation_firstpage" content="149"/> <meta name="citation_lastpage" content="165"/> <meta name="citation_article_type" content="Article"/> <meta name="citation_fulltext_world_readable" content=""/> <meta name="citation_language" content="en"/> <meta name="dc.identifier" content="doi:10.1007/s10539-014-9469-4"/> <meta name="DOI" content="10.1007/s10539-014-9469-4"/> <meta name="size" content="219747"/> <meta name="citation_doi" content="10.1007/s10539-014-9469-4"/> <meta name="citation_springer_api_url" content="http://api.springer.com/xmldata/jats?q=doi:10.1007/s10539-014-9469-4&api_key="/> <meta name="description" content="Phenomenal consciousness or the subjective experience of feeling sensory stimuli is fundamental to human existence. Because of the ubiquity of their subjec"/> <meta name="dc.creator" content="Key, Brian"/> <meta name="dc.subject" content="Philosophy of Biology"/> <meta name="dc.subject" content="Evolutionary Biology"/> <meta name="citation_reference" content="citation_journal_title=Epilepsia; citation_title=Anatomofunctional organization of the insular cortex: a study using intracerebral electrical stimulation in epileptic patients; citation_author=A Afif, L Minotti, P Kahane, D Hoffman; citation_volume=51; citation_publication_date=2010; citation_pages=2305-2315; citation_id=CR1"/> <meta name="citation_reference" content="citation_journal_title=PNAS; citation_title=Spatioptemporal dynamics in understanding hand-object interactions; citation_author=P Avanzini, M Fabbri-Destro, C Campi, A Pascarella, G Barchiesi, L Cattaneo, G Rizzolatti; citation_volume=110; citation_publication_date=2013; citation_pages=15878-15885; citation_id=CR2"/> <meta name="citation_reference" content="citation_journal_title=Front Psychol; citation_title=Global workspace dynamics: cortical “binding and propagation” enables conscious contents; citation_author=BJ Baars, S Franklin, TZ Ramsoy; citation_publication_date=2013; citation_id=CR3"/> <meta name="citation_reference" content="citation_journal_title=Pain; citation_title=Spinal-, brainstem- and cerebrally mediated responses at- and below-level of a spinal cord contusion in rats: evaluation of pain-like behavior; citation_author=C Baastrup, CC Maersk-Moller, JR Nyengaard, TS Jensen, NB Finnerup; citation_volume=151; citation_publication_date=2010; citation_pages=670-679; citation_id=CR4"/> <meta name="citation_reference" content="citation_journal_title=Neuron; citation_title=Canonical microcircuits for predictive coding; citation_author=AM Bastos, WM Usrey, RA Adams, GR Mangun, P Fries, KJ Friston; citation_volume=76; citation_publication_date=2012; citation_pages=695-711; citation_id=CR5"/> <meta name="citation_reference" content="citation_journal_title=J Neurophysiol; citation_title=Multiple somatotopic representations of heat and mechanical pain in the operculo-insular cortex: a high-resolution fMRI study; citation_author=U Baumgartner, GD Iannetti, L Zambreanu, P Stoeter, R-D Treede, I Tracey; citation_volume=104; citation_publication_date=2010; citation_pages=2863-2872; citation_id=CR150"/> <meta name="citation_reference" content="citation_journal_title=Curr Opin Neurobiol; citation_title=Maps in birds: representational mechanisms and neural bases; citation_author=VP Bingman, KP Able; citation_volume=12; citation_publication_date=2002; citation_pages=745-750; citation_id=CR7"/> <meta name="citation_reference" content="citation_journal_title=Behav Brain Sci; citation_title=A perceptual-defensive-recuperative model of fear and pain; citation_author=RC Bolles, MS Faneslow; citation_volume=3; citation_publication_date=1980; citation_pages=291-323; citation_id=CR8"/> <meta name="citation_reference" content="citation_journal_title=Front Psychol; citation_title=Epilepsy as a neurodevelopmental disorder; citation_author=Y Bozzi, S Casarosa, M Caleo; citation_publication_date=2012; citation_id=CR9"/> <meta name="citation_reference" content="citation_journal_title=Brain Behav Evol; citation_title=Stalking the everted telencephalon: comparisons of forebrain organization in basal ray-finned fishes and teleosts; citation_author=MR Braford; citation_volume=74; citation_publication_date=2009; citation_pages=56-76; citation_id=CR10"/> <meta name="citation_reference" content="citation_journal_title=Dis Aquat Org; citation_title=Pain perception, aversion and fear in fish; citation_author=VA Braithwaite, P Boulcott; citation_volume=75; citation_publication_date=2007; citation_pages=131-138; citation_id=CR11"/> <meta name="citation_reference" content="citation_journal_title=J Agric Environ Ethics; citation_title=Variation in emotion and cognition in fish; citation_author=VA Braithwaite, F Huntingford, R Bos; citation_volume=26; citation_publication_date=2013; citation_pages=7-23; citation_id=CR12"/> <meta name="citation_reference" content="citation_journal_title=Dis Aquat Org; citation_title=Welfare in aquatic organisms—is there some faith-based HAR-King going on here?; citation_author=HI Browman, AB Skiftesvik; citation_volume=94; citation_publication_date=2011; citation_pages=255-257; citation_id=CR13"/> <meta name="citation_reference" content="citation_journal_title=Anat Rec; citation_title=Chordate evolution and the origin of the craniates: an old brain in a new head; citation_author=AB Butler; citation_volume=261; citation_publication_date=2000; citation_pages=111-125; citation_id=CR14"/> <meta name="citation_reference" content="citation_journal_title=Brain; citation_title=The minicolumn hypothesis in neuroscience; citation_author=DP Buxhoeveden, MF Casanova; citation_volume=125; citation_publication_date=2002; citation_pages=935-951; citation_id=CR15"/> <meta name="citation_reference" content="citation_journal_title=Encycl Behav Neurosci; citation_title=Active avoidance and escape learning; citation_author=CK Cain, J-S Choi, JE LeDoux; citation_volume=1; citation_publication_date=2010; citation_pages=1-9; citation_id=CR16"/> <meta name="citation_reference" content="citation_journal_title=Transl Neurosci; citation_title=Cortical organization: anatomical findings based on systems theory; citation_author=MF Casanova; citation_volume=1; citation_publication_date=2010; citation_pages=62-71; citation_id=CR17"/> <meta name="citation_reference" content="citation_journal_title=Appl Anim Behav Sci; citation_title=Can fish suffer? Perspectives on sentience, pain, fear and stress; citation_author=KP Chandroo, IJH Duncan, RD Moccia; citation_volume=86; citation_publication_date=2004; citation_pages=225-250; citation_id=CR18"/> <meta name="citation_reference" content="citation_journal_title=J Neurophysiol; citation_title=Fine-scale organization of SI (area 3b) in the squirrel monkey revealed with intrinsic optical imaging; citation_author=LM Chen, RM Friedman, BM Ramsden, RH LaMotte, AW Roe; citation_volume=86; citation_publication_date=2001; citation_pages=3011-3029; citation_id=CR19"/> <meta name="citation_reference" content="citation_journal_title=Pain; citation_title=High-resolution functional magnetic resonance imaging of noxious heat and activations along the central sulcus in New World monkeys; citation_author=LM Chen, BC Dillenburger, F Wang, RM Friedman, MJ Avison; citation_volume=152; citation_publication_date=2011; citation_pages=522-532; citation_id=CR20"/> <meta name="citation_reference" content="citation_journal_title=J Ichthyol; citation_title=Pain sensitivity of behavior of fishes; citation_author=LS Chervova; citation_volume=37; citation_publication_date=1997; citation_pages=98-102; citation_id=CR21"/> <meta name="citation_reference" content="citation_journal_title=Behav Biol; citation_title=Telencephalic lesions and behavior in the teleost, macropodus opercularis: reproductive, startle reaction, and operant behavior in the male; citation_author=RE Davis, J Kassel, P Schwagmeyer; citation_volume=18; citation_publication_date=1976; citation_pages=165-177; citation_id=CR22"/> <meta name="citation_reference" content="citation_journal_title=Brain Behav Evol; citation_title=The pallium and mind/behavior relationships in teleost fishes; citation_author=LS Demski; citation_volume=82; citation_publication_date=2013; citation_pages=31-44; citation_id=CR23"/> <meta name="citation_reference" content="citation_journal_title=Brain; citation_title=Studies of spasticity in man. 3. Analysis of reflex activity evoked by noxious cutaneous stimulation; citation_author=MR Dimitrijevic, PW Nathan; citation_volume=91; citation_publication_date=1968; citation_pages=349-368; citation_id=CR24"/> <meta name="citation_reference" content="citation_journal_title=PNAS; citation_title=Cell-type homologies and the origins of the neocortex; citation_author=J Dugas-Ford, JJ Rowell, CW Ragsdale; citation_volume=109; citation_publication_date=2012; citation_pages=16974-16979; citation_id=CR25"/> <meta name="citation_reference" content="citation_journal_title=J Pain; citation_title=Mechanoreceptive and nociceptive responses in the central nervous system of goldfish (Carassius auratus) and trout (Oncorrhynchus mykiss); citation_author=R Dunlop, P Laming; citation_volume=6; citation_publication_date=2005; citation_pages=561-568; citation_id=CR26"/> <meta name="citation_reference" content="citation_journal_title=Appl Anim Behav Sci; citation_title=Avoidance learning in goldfish (Carassius auratus) and implications for pain perception; citation_author=R Dunlop, S Millsopp, P Laming; citation_volume=976; citation_publication_date=2006; citation_pages=255-271; citation_id=CR27"/> <meta name="citation_reference" content="citation_journal_title=Science; citation_title=Forebrain connections in the goldfish support telencephalic homologies with land vertebrates; citation_author=SM Echleter, WM Saidel; citation_volume=212; citation_publication_date=1981; citation_pages=683-685; citation_id=CR28"/> <meta name="citation_reference" content="citation_title=Fin erosion in farmed fish; citation_inbook_title=Fish welfare, chapter 9; citation_publication_date=2008; citation_id=CR29; citation_author=T Ellis; citation_author=B Oidtmann; citation_author=S St Hilaire; citation_author=JF Turnbull; citation_author=BP North; citation_author=CM MacIntyre; citation_author=J Nikolaidis; citation_author=I Hoyle; citation_author=SC Kestin; citation_author=TG Knowles; citation_publisher=Wiley"/> <meta name="citation_reference" content="citation_journal_title=Fish Physiol Biochem; citation_title=Diel changes in plasma cortisol and effects of size and stress duration on the cortisol response in European sea bass (Dicentrarchus labrax); citation_author=E Fatira, N Papandroulakis, M Pavlidis; citation_publication_date=2014; citation_id=CR30"/> <meta name="citation_reference" content="citation_journal_title=Gen Comp Endocrinol; citation_title=CRF and stress in fish; citation_author=G Filk, PHM Klaren, EH Burg, JR Metz, MO Huising; citation_volume=146; citation_publication_date=2006; citation_pages=36-44; citation_id=CR31"/> <meta name="citation_reference" content="citation_journal_title=Physiol Behav; citation_title=Teleost telencephalon and learning: an interpretive review of data and hypotheses; citation_author=NC Flood, JB Overmier, GE Savage; citation_volume=16; citation_publication_date=1976; citation_pages=783-798; citation_id=CR32"/> <meta name="citation_reference" content="citation_journal_title=Comp Biochem Physiol A: Mol Integr Physiol; citation_title=The effects of caudal fin amputation on metabolic interaction between digestion and locomotion in juveniles of three cyprinid fish species with different metabolic modes; citation_author=C Fu, ZD Cao, SJ Fu; citation_volume=164; citation_publication_date=2013; citation_pages=456-465; citation_id=CR33"/> <meta name="citation_reference" content="citation_journal_title=J Comp Neurol; citation_title=Organization of the gymnotiform fish pallium in relation to learning and memory: III. Intrinsic connections; citation_author=ACC Giassi, W Ellis, L Maler; citation_volume=520; citation_publication_date=2012; citation_pages=3369-3394; citation_id=CR34"/> <meta name="citation_reference" content="citation_journal_title=Neurobiol Dis; citation_title=Neuronal migration disorders; citation_author=R Guerrini, E Parrini; citation_volume=38; citation_publication_date=2010; citation_pages=154-166; citation_id=CR35"/> <meta name="citation_reference" content="citation_journal_title=TINS; citation_title=Abnormal development of the human cerebral cortex: genetics, functional consequences and treatment options; citation_author=R Guerrini, WB Dobyns, AJ Barkovich; citation_volume=31; citation_publication_date=2008; citation_pages=154-162; citation_id=CR36"/> <meta name="citation_reference" content="citation_journal_title=Curr Opin Neurobiol; citation_title=The avian ‘prefrontal cortex’ and cognition; citation_author=O Gunturkun; citation_volume=15; citation_publication_date=2005; citation_pages=686-693; citation_id=CR37"/> <meta name="citation_reference" content="citation_journal_title=Psychol Res; citation_title=The convergent evolution of neural substrates for cognition; citation_author=O Gunturkun; citation_volume=76; citation_publication_date=2012; citation_pages=212-219; citation_id=CR38"/> <meta name="citation_reference" content="citation_journal_title=J Comp Physiol Psychol; citation_title=Specific and permanent deficits in instrumental avoidance responding following forebrain ablation in the goldfish; citation_author=FR Hainsworth, JB Overmier, CT Snowdon; citation_volume=63; citation_publication_date=1967; citation_pages=111-116; citation_id=CR39"/> <meta name="citation_reference" content="citation_journal_title=Dev Biol; citation_title=Are pioneer axons guided by regulatory gene expression domains in the zebrafish forebrain? High-resolution analysis of the patterning of the zebrafish brain during axon tract formation; citation_author=JT Hjorth, B Key; citation_volume=229; citation_publication_date=2001; citation_pages=271-286; citation_id=CR40"/> <meta name="citation_reference" content="citation_journal_title=Univ Psychol; citation_title=Neuronal mechanisms of learning in teleost fish; citation_author=C Hurtado-Parrado; citation_volume=9; citation_publication_date=2010; citation_pages=663-672; citation_id=CR41"/> <meta name="citation_reference" content="citation_journal_title=Brain Behav Evol; citation_title=Evolutionary perspectives on the function of the optic tectum; citation_author=D Ingle; citation_volume=8; citation_publication_date=1973; citation_pages=211-237; citation_id=CR42"/> <meta name="citation_reference" content="citation_journal_title=PLoS ONE; citation_title=The temporal dynamics of early visual cortex involvement in behavioral priming; citation_author=C Jacobs, TA Gaff, R Goebel, AT Sack; citation_volume=7; citation_publication_date=2012; citation_pages=e48808; citation_id=CR43"/> <meta name="citation_reference" content="citation_journal_title=J Comp Neurol; citation_title=Global view of the functional molecular organization of the avian cerebrum: mirror images and functional columns; citation_author=ED Jarvis, J Yu, MV Rivas, H Horita, G Feenders, O Whitney, SC Jarvis, ER Jarvis, L Kubikova, AEP Puck, C Siang-Bakshi, S Martin, M McElroy, E Hara, J Howard, A Pfenning, H Mouritsen, C-C Chen, K Wada; citation_volume=521; citation_publication_date=2013; citation_pages=3614-3665; citation_id=CR44"/> <meta name="citation_reference" content="citation_journal_title=Biol Philos; citation_title=Science, sentience, and animal welfare; citation_author=RC Jones; citation_volume=28; citation_publication_date=2013; citation_pages=1-30; citation_id=CR45"/> <meta name="citation_reference" content="citation_journal_title=Lancet; citation_title=Sites of action of morphine in the brain; citation_author=AKP Jones, KJ Friston, LY Qi, M Harris, VJ Cunningham, T Jones, C Feinman, RSJ Frackowiak; citation_volume=338; citation_publication_date=1991; citation_pages=825; citation_id=CR46"/> <meta name="citation_reference" content="citation_journal_title=J Vet Pharmacol Ther; citation_title=The dose-response relation for the antinociceptive effect of morphine in a fish, rainbow trout; citation_author=SG Jones, C Kamunde, K Lemke, ED Stevens; citation_volume=35; citation_publication_date=2012; citation_pages=563-570; citation_id=CR47"/> <meta name="citation_reference" content="citation_journal_title=Brain Res Bull; citation_title=Topographic maps are fundamental to sensory processing; citation_author=JH Kaas; citation_volume=44; citation_publication_date=1997; citation_pages=107-112; citation_id=CR48"/> <meta name="citation_reference" content="citation_journal_title=PNAS; citation_title=Evolution of columns, modules, and domains in the neocortex of primates; citation_author=JH Kaas; citation_volume=109; citation_publication_date=2012; citation_pages=10655-10660; citation_id=CR49"/> <meta name="citation_reference" content="citation_journal_title=Curr Biol; citation_title=Qualia; citation_author=R Kanai, N Tsuchiya; citation_volume=22; citation_publication_date=2012; citation_pages=R392-R396; citation_id=CR50"/> <meta name="citation_reference" content="citation_journal_title=Anim Behav; citation_title=Effect of forebrain ablation on the performance of a conditioned avoidance response in the teleost fish, Tilapia H. Macrocephala; citation_author=H Kaplan, LR Aronson; citation_volume=15; citation_publication_date=1967; citation_pages=438-448; citation_id=CR51"/> <meta name="citation_reference" content="citation_journal_title=Brain Res Bull; citation_title=Evolutionary and comparative aspects of nociception; citation_author=M Kavaliers; citation_volume=21; citation_publication_date=1988; citation_pages=923-931; citation_id=CR52"/> <meta name="citation_reference" content="citation_journal_title=Conscious Cogn; citation_title=Insight without cortex: lessons from the avian brain; citation_author=JA Kirsch, O Gunturkun, J Rose; citation_volume=17; citation_publication_date=2008; citation_pages=475-483; citation_id=CR53"/> <meta name="citation_reference" content="Kittilsen S (2013) Functional aspects of emotions in fish. Behav Process 100:153–159"/> <meta name="citation_reference" content="citation_journal_title=Neuroimage; citation_title=Visual feature binding: the critical time windows of V1/V2 and parietal activity; citation_author=M Koivisto, J Silvanto; citation_volume=59; citation_publication_date=2012; citation_pages=1608-1614; citation_id=CR54"/> <meta name="citation_reference" content="citation_journal_title=Neuroimage; citation_title=The role of early visual corex (V1/V2) in conscious and unconscious visual perception; citation_author=M Koivisto, T Mantyla, J Silvanto; citation_volume=51; citation_publication_date=2010; citation_pages=828-834; citation_id=CR55"/> <meta name="citation_reference" content="citation_journal_title=Conscious Cogn; citation_title=Transcranial magnetic stimulation of early visual cortex interferes with subjective visual awareness and objective forced-choice performance; citation_author=M Koivisto, H Railo, N Salminen-Vaparanta; citation_volume=20; citation_publication_date=2011; citation_pages=288-298; citation_id=CR56"/> <meta name="citation_reference" content="citation_journal_title=Curr Opin Neurobiol; citation_title=Neural maps in the electrosensory system of weakly electric fish; citation_author=R Krahe, L Maler; citation_volume=24; citation_publication_date=2014; citation_pages=13-21; citation_id=CR57"/> <meta name="citation_reference" content="citation_journal_title=Neurosci Biobehav Rev; citation_title=Compensatory plasticity and cross-modal reorganization following early visual deprivation; citation_author=R Kupers, M Pitto; citation_publication_date=2013; citation_id=CR58"/> <meta name="citation_reference" content="citation_journal_title=TINS; citation_title=A cellular mechanism for cortical associations: an organizing principle for the cerebral cortex; citation_author=M Larkum; citation_volume=36; citation_publication_date=2013; citation_pages=141-151; citation_id=CR59"/> <meta name="citation_reference" content="citation_journal_title=Brain Res Rev; citation_title=Involvement of the optic tectum and mesencephalic reticular formation in the generation of saccadic eye movements in goldfish; citation_author=MA Luque, MP Perez-Perez, L Herrero, B Torres; citation_volume=49; citation_publication_date=2005; citation_pages=388-397; citation_id=CR60"/> <meta name="citation_reference" content="citation_journal_title=Front Syst Neurosci; citation_title=Distinct superficial and deep laminar domains of activity in the visual cortex during rest and stimulation; citation_author=A Maier, GK Adams, C Aura, DA Leopold; citation_publication_date=2010; citation_id=CR61"/> <meta name="citation_reference" content="citation_journal_title=J Cell Physiol; citation_title=The zebrafish as a model for nociception studies; citation_author=V Malafoglia, B Bryant, W Raffaeli, A Giordano, G Bellipanni; citation_volume=228; citation_publication_date=2013; citation_pages=1956-1966; citation_id=CR62"/> <meta name="citation_reference" content="citation_journal_title=J Neurosci; citation_title=Fine-grained nociceptive maps in primary somatosensory cortex; citation_author=F Mancini, P Haggard, GD Iannetti, MR Longo, MI Sereno; citation_volume=32; citation_publication_date=2012; citation_pages=17155-17162; citation_id=CR63"/> <meta name="citation_reference" content="citation_journal_title=Sci Total Environ; citation_title=The interactive effects of multiple stressors on physiological stress responders and club cell investment in fathead minnows; citation_author=AK Manek, MC Ferrari, S Niyogi, DP Chivers; citation_volume=467–477; citation_publication_date=2014; citation_pages=90-97; citation_id=CR64"/> <meta name="citation_reference" content="citation_journal_title=Eur J Neurosci; citation_title=Multiple maps and activity-dependent representational plasticity in the anterior Wust of the adult barn owl (Tyto alba); citation_author=PR Manger, GN Elston, JD Pettigrew; citation_volume=16; citation_publication_date=2002; citation_pages=743-750; citation_id=CR65"/> <meta name="citation_reference" content="citation_journal_title=Pain; citation_title=Formalin pain is expressed in decerebrate rats but not attenuated by morphine; citation_author=BK Matthies, KBJ Franklin; citation_volume=51; citation_publication_date=1992; citation_pages=199-206; citation_id=CR66"/> <meta name="citation_reference" content="citation_journal_title=Behav Brain Res; citation_title=Effects of partial decortication on the opioid analgesia in the formalin test; citation_author=BK Matthies, KBJ Franklin; citation_volume=67; citation_publication_date=1995; citation_pages=59-66; citation_id=CR67"/> <meta name="citation_reference" content="citation_journal_title=Neuroimage; citation_title=Spatial segregation of somatosensory and pain activations in the human operculo-insular cortex; citation_author=L Mazzola, I Faillenot, F-G Barral, F Mauguiere, R Peyron; citation_volume=60; citation_publication_date=2012; citation_pages=5409-5418; citation_id=CR68"/> <meta name="citation_reference" content="citation_journal_title=Brain Res; citation_title=Functional anatomy of the tectum mesencephala of the goldfish. An explorative analysis of the functional implications of the laminar structural organization of the tectum; citation_author=J Meek; citation_volume=287; citation_publication_date=1983; citation_pages=247-297; citation_id=CR69"/> <meta name="citation_reference" content="citation_journal_title=Brain; citation_title=The columnar organization of the neocortex; citation_author=VB Mountcastle; citation_volume=120; citation_publication_date=1997; citation_pages=701-722; citation_id=CR70"/> <meta name="citation_reference" content="citation_journal_title=Brain Behav Evol; citation_title=An evolutionary interpretation of teleostean forebrain anatomy; citation_author=T Mueller, MF Wullimann; citation_volume=74; citation_publication_date=2009; citation_pages=30-42; citation_id=CR71"/> <meta name="citation_reference" content="citation_journal_title=Brain Res; citation_title=The dorsal pallium in zebrafish, Danio rerio (Cyprinidae, Teleostei); citation_author=T Mueller, Z Dong, MA Berberoglu, S Guo; citation_volume=1381; citation_publication_date=2011; citation_pages=95-105; citation_id=CR72"/> <meta name="citation_reference" content="citation_journal_title=J Neurosci; citation_title=A dissociation of motion and spatial-pattern vision in the avian telencephalon: implications for the evolution of “visual streams”; citation_author=AP Nguyen, ML Spetch, NA Crowder, IR Winship, PL Hurd, DR Wylie; citation_volume=24; citation_publication_date=2004; citation_pages=4962-4970; citation_id=CR73"/> <meta name="citation_reference" content="citation_journal_title=Anat Embryol; citation_title=The neocortex. An overview of its evolutionary development, structural organization and synaptology; citation_author=R Nieuwenhuys; citation_volume=190; citation_publication_date=1994; citation_pages=307-337; citation_id=CR74"/> <meta name="citation_reference" content="citation_journal_title=J Comp Physiol A; citation_title=Somatosensory evoked potentials in the telencephalon of Atlanic salmon (Salmo salar) following galvanic stimulation of the tail; citation_author=J Nordgreen, TE Horsberg, B Ranheim, ACN Chen; citation_volume=193; citation_publication_date=2007; citation_pages=1235-1242; citation_id=CR75"/> <meta name="citation_reference" content="citation_journal_title=Brain Res Bull; citation_title=Forebrain evolution in bony fishes; citation_author=RG Northcutt; citation_volume=75; citation_publication_date=2008; citation_pages=191-205; citation_id=CR76"/> <meta name="citation_reference" content="citation_journal_title=Brain Behav Evol; citation_title=Do teleost fishes possess a homolog of mammalian isocortex?; citation_author=RG Northcutt; citation_volume=78; citation_publication_date=2011; citation_pages=136-138; citation_id=CR77"/> <meta name="citation_reference" content="citation_journal_title=Brain Behav Evol; citation_title=A pallial visual area in the telencephalon of the bony fish Polypterus ; citation_author=RG Northcutt, W Plassman, PH Holmes, WM Saidel; citation_volume=64; citation_publication_date=2004; citation_pages=1-10; citation_id=CR78"/> <meta name="citation_reference" content="citation_journal_title=Nature; citation_title=Sparse coding and high-order correlations in fine-scale cortical networks; citation_author=IE Ohiorhenuan, F Mechler, KP Purpura, AM Schmid, Q Hu, JD Victor; citation_volume=466; citation_publication_date=2010; citation_pages=617-622; citation_id=CR79"/> <meta name="citation_reference" content="citation_journal_title=Front Syst Neurosci; citation_title=Inter-laminar microcircuits across neocortex: repair and augmentation; citation_author=I Opris; citation_volume=7; citation_publication_date=2013; citation_pages=80; citation_id=CR80"/> <meta name="citation_reference" content="citation_journal_title=Front Neural Circuits; citation_title=Closing the loop in primate prefrontal cortex: inter-laminar processing; citation_author=I Opris, JL Fuqua, PF Huetti, GA Gerhardt, TW Berger, RE Hampson, SA Deadwyler; citation_volume=6; citation_publication_date=2012; citation_pages=1-13; citation_id=CR81"/> <meta name="citation_reference" content="citation_journal_title=Behav Biol; citation_title=Effects of telencephalic ablation upon nest-building and avoidance bahaviors in East African mouthbreeding fish, Tilapia mossambica ; citation_author=JB Overmier, D Gross; citation_volume=12; citation_publication_date=1974; citation_pages=211-222; citation_id=CR82"/> <meta name="citation_reference" content="citation_journal_title=Behav Neurosci; citation_title=Serial ablations of the telencephalon and avoidance learning by goldfish (Carassius auratus); citation_author=JB Overmier, MR Papini; citation_volume=99; citation_publication_date=1985; citation_pages=509-520; citation_id=CR83"/> <meta name="citation_reference" content="citation_journal_title=Exp Brain Res; citation_title=Connectivity of the goldfish optic tectum with the mesencephalic and rhombencephalic reticular formation; citation_author=MP Perez-Perez, MA Luque, L Herrero, PA Nunez-Abades, B Torres; citation_volume=151; citation_publication_date=2003; citation_pages=123-135; citation_id=CR84"/> <meta name="citation_reference" content="citation_journal_title=Pharmacol Biochem Behav; citation_title=Localization of the antinociceptive action of morphine in primate brain; citation_author=A Pert, T Yaksh; citation_volume=3; citation_publication_date=1975; citation_pages=133-138; citation_id=CR85"/> <meta name="citation_reference" content="citation_journal_title=Dev Neurobiol; citation_title=Glutamate drives the touch response through a rostral loop in the spinal cord of zebrafish embryos; citation_author=T Pietri, E Manalo, J Ryan, L Saint-Amant, P Washbourne; citation_volume=69; citation_publication_date=2009; citation_pages=780-795; citation_id=CR86"/> <meta name="citation_reference" content="citation_journal_title=Cereb Cortex; citation_title=On the emergence of primary visual perception; citation_author=DA Pollen; citation_volume=21; citation_publication_date=2011; citation_pages=1941-1953; citation_id=CR87"/> <meta name="citation_reference" content="citation_journal_title=Eur J Neurosci; citation_title=Emotional and spatial learning in goldfish is dependent on different telencephalic pallial systems; citation_author=M Portavella, JP Vargas; citation_volume=21; citation_publication_date=2005; citation_pages=2800-2806; citation_id=CR88"/> <meta name="citation_reference" content="citation_journal_title=Physiol Behav; citation_title=Involvement of the telencephalon in spaced-trial avoidance learning in the goldfish (Carassius auratus); citation_author=M Portavella, CM Salas, JP Vargas, MR Papini; citation_volume=80; citation_publication_date=2003; citation_pages=49-56; citation_id=CR89"/> <meta name="citation_reference" content="citation_journal_title=J Neurosci; citation_title=Avoidance response in goldfish: emotional and temporal involvement of medial and lateral telencephalic pallium; citation_author=M Portavella, B Torres, C Salas; citation_volume=24; citation_publication_date=2004; citation_pages=2335-2342; citation_id=CR90"/> <meta name="citation_reference" content="citation_journal_title=Neurosci Lett; citation_title=Lesions of the medial pallium, but not of the lateral pallium, disrupt spaced-trial avoidance learning in goldfish (Carassius auratus); citation_author=M Portavella, B Torres, C Salas, MR Papini; citation_volume=362; citation_publication_date=2004; citation_pages=75-78; citation_id=CR91"/> <meta name="citation_reference" content="citation_journal_title=J Neurosci; citation_title=Sensory processing in the pallium of a Mormyrid fish; citation_author=JC Precht, G Emde, J Wolfart, S Karamursel, GN Akoev, YN Andrianov, TH Bullock; citation_volume=18; citation_publication_date=1998; citation_pages=7381-7393; citation_id=CR92"/> <meta name="citation_reference" content="citation_title=Quantitative trait loci for magnitude of the plasma cortisol response to confinement in rainbow trout; citation_publication_date=2014; citation_id=CR93; citation_author=E Quillet; citation_author=F Krieg; citation_author=N Dechamp; citation_author=C Hervet; citation_author=A Bérard; citation_author=P Roy; citation_author=R Guyomard; citation_author=P Prunet; citation_author=TG Pottinger; citation_publisher=Genet"/> <meta name="citation_reference" content="citation_journal_title=Cortex; citation_title=Two means of suppressing visual awareness: a direct comparison of visual masking and transcranial magnetic stimulation; citation_author=H Railo, M Koivisto; citation_volume=48; citation_publication_date=2012; citation_pages=333-343; citation_id=CR94"/> <meta name="citation_reference" content="citation_journal_title=Appl Anim Behav Sci; citation_title=Behavioural analysis of a nociceptive event in fish: comparisons between three species demonstrate specific responses; citation_author=SC Reilly, JP Quinn, AR Cossins, LU Sneddon; citation_volume=114; citation_publication_date=2008; citation_pages=248-259; citation_id=CR95"/> <meta name="citation_reference" content="citation_journal_title=J Neurosci; citation_title=Visuomotor behaviors in larval zebrafish after GFP-guided laser ablation of the optic tectum; citation_author=T Roeser, H Naier; citation_volume=23; citation_publication_date=2003; citation_pages=3726-3734; citation_id=CR96"/> <meta name="citation_reference" content="citation_journal_title=Physiol Behav; citation_title=Tailfin clipping, a painful procedure: studies on Nile tilapia and common carp; citation_author=JAC Roques, W Abbink, F Geurds, H Vis, G Flik; citation_volume=101; citation_publication_date=2010; citation_pages=533-540; citation_id=CR97"/> <meta name="citation_reference" content="citation_journal_title=Rev Fish Sci; citation_title=The neurobiological nature of fishes and the question of awareness and pain; citation_author=JD Rose; citation_volume=10; citation_publication_date=2002; citation_pages=1-38; citation_id=CR98"/> <meta name="citation_reference" content="citation_journal_title=Dis Aquat Organ; citation_title=Anthropomorphism and ‘mental welfare’ of fishes; citation_author=JD Rose; citation_volume=75; citation_publication_date=2007; citation_pages=139-154; citation_id=CR99"/> <meta name="citation_reference" content="citation_journal_title=Fish Fish; citation_title=Can fish really feel pain?; citation_author=JD Rose, R Arlinghaus, SJ Cooke, BK Diggles, W Sawynok, ED Stevens, CDL Wynne; citation_volume=15; citation_publication_date=2014; citation_pages=97-133; citation_id=CR100"/> <meta name="citation_reference" content="citation_journal_title=Brain Res; citation_title=Identification of visual pallial telencephalon in the goldfish, Carassius auratus: a combined cytochrome oxidase and electrophysiological study; citation_author=WM Saidel, K Marquez-Houston, AB Butler; citation_volume=919; citation_publication_date=2001; citation_pages=82-93; citation_id=CR101"/> <meta name="citation_reference" content="citation_journal_title=Zebrafish; citation_title=Development of motor networks in zebrafish embryos; citation_author=L Saint-Amant; citation_volume=3; citation_publication_date=2006; citation_pages=173-190; citation_id=CR102"/> <meta name="citation_reference" content="citation_journal_title=Nature; citation_title=Temporal factors in avoidance learning in normal and forebrainless goldfish (Cartassius auratus); citation_author=GE Savage; citation_volume=218; citation_publication_date=1968; citation_pages=1168-1169; citation_id=CR103"/> <meta name="citation_reference" content="citation_journal_title=Anim Behav; citation_title=Some preliminary observations on the role of the telencephalon in food-reinforced behaviour in the goldfish, Carassius auratus ; citation_author=GE Savage; citation_volume=17; citation_publication_date=1969; citation_pages=760-772; citation_id=CR104"/> <meta name="citation_reference" content="citation_journal_title=Brain Struct Funct; citation_title=Mapping functional connectivity in barrel-related columns reveals layer- and cell type-specific microcircuits; citation_author=D Schubert, R Kotter, JF Staiger; citation_volume=212; citation_publication_date=2007; citation_pages=107-119; citation_id=CR105"/> <meta name="citation_reference" content="citation_journal_title=Vet Clin Exot Anim; citation_title=Fish analgesia: pain, stress, fear aversion or nociception?; citation_author=E Scott Weber; citation_volume=14; citation_publication_date=2011; citation_pages=21-32; citation_id=CR106"/> <meta name="citation_reference" content="citation_journal_title=Comp Cogn Behav Rev; citation_title=Why can birds be so smart? Background, significance, and implications of the revised view of the avian brain; citation_author=T Shimizu; citation_volume=4; citation_publication_date=2009; citation_pages=103-115; citation_id=CR107"/> <meta name="citation_reference" content="citation_journal_title=Behav Brain Res; citation_title=Visual circuits of the avian telencephalon; citation_author=T Shimizu, AN Bowers; citation_volume=98; citation_publication_date=1999; citation_pages=183-191; citation_id=CR108"/> <meta name="citation_reference" content="citation_journal_title=J Comp Neurol; citation_title=Intratelencephalic projections of the visual wulst in pigeons (Columba livia); citation_author=T Shimizu, K Cox, HJ Karten; citation_volume=359; citation_publication_date=1995; citation_pages=551-572; citation_id=CR109"/> <meta name="citation_reference" content="citation_journal_title=TINS; citation_title=Always returning: feedback and sensory processing in visual cortex and thalamus; citation_author=AM Sillito, J Cudeiro, HE Jones; citation_volume=29; citation_publication_date=2006; citation_pages=307-316; citation_id=CR110"/> <meta name="citation_reference" content="citation_journal_title=Appl Anim Behave Sci; citation_title=The evidence for pain in fish: the use of morphine as an analgesic; citation_author=LU Sneddon; citation_volume=83; citation_publication_date=2003; citation_pages=153-162; citation_id=CR111"/> <meta name="citation_reference" content="citation_journal_title=Brain Res Rev; citation_title=Evolution of nociception in vertebrates: comparative analysis of lower vertebrates; citation_author=LU Sneddon; citation_volume=46; citation_publication_date=2004; citation_pages=123-130; citation_id=CR112"/> <meta name="citation_reference" content="citation_title=Nociception or pain in fish; citation_inbook_title=Encyclopedia of fish physiology; citation_publication_date=2011; citation_pages=713-719; citation_id=CR113; citation_author=LU Sneddon; citation_publisher=Academic Press"/> <meta name="citation_reference" content="citation_journal_title=Proc R Soc Lond B; citation_title=Do fishes have nociceptors? Evidence for the evolution of a vertebrate sensory system; citation_author=LU Sneddon, VA Braithwaite, MJ Gentle; citation_volume=270; citation_publication_date=2003; citation_pages=1115-1121; citation_id=CR114"/> <meta name="citation_reference" content="citation_journal_title=J Neurosci Res; citation_title=Topical application of morphine to the rat somatosensory cortex produces analgesia to tonic pain; citation_author=R Soto-Moyano, J Galvez, C Vallejos, A Hernandez; citation_volume=19; citation_publication_date=1988; citation_pages=511-514; citation_id=CR115"/> <meta name="citation_reference" content="citation_journal_title=Science; citation_title=Reversal of interlaminar signal between sensory and memory processing in monkey temporal cortex; citation_author=D Takeuchi, T Hirabayashi, K Tamura, Y Miyashita; citation_volume=331; citation_publication_date=2011; citation_pages=1443-1447; citation_id=CR116"/> <meta name="citation_reference" content="citation_journal_title=Neuropsychopharmacology; citation_title=Naloxone-reversible modulation of pain circuitry by left prefrontal rTMS; citation_author=JJ Taylor, JJ Borckardt, M Canterberyy, X Li, CA Hanlon, TR Brown, MS George; citation_volume=38; citation_publication_date=2013; citation_pages=1189-1197; citation_id=CR117"/> <meta name="citation_reference" content="citation_journal_title=TINS; citation_title=The topographic brain: from neural connectivity to cognition; citation_author=J-P Thivierge, GF Marcus; citation_volume=30; citation_publication_date=2007; citation_pages=251-258; citation_id=CR118"/> <meta name="citation_reference" content="citation_journal_title=J Neurophysiol; citation_title=Anterior parietal cortical response to tactile and skin-heating stimuli applied to the same skin site; citation_author=M Tommerdahl, KA Delemos, CJ Vierck, OV Favorov, BL Whitsel; citation_volume=75; citation_publication_date=1996; citation_pages=2662-2670; citation_id=CR119"/> <meta name="citation_reference" content="citation_journal_title=Neuroimage; citation_title=Nociceptive and non-nociceptive sub-regions in the human somatosensory cortex: an MEG study using fMRI constraints; citation_author=K Torquati, V Pizzella, C Babiloni, C Gratta, S Della Penna, A Ferretti, R Franciotti, PM Rossini, GL Romani; citation_volume=26; citation_publication_date=2005; citation_pages=48-56; citation_id=CR120"/> <meta name="citation_reference" content="citation_journal_title=J Neurosci; citation_title=Selective cortical layering abnormalities and behavioral deficits in cortex-specific Pax6 knock-out mice; citation_author=TC Tuoc, K Radyushkin, AB Tonchev, MC Piñon, R Ashery-Padan, Z Molnár, MS Davidoff, A Stoykova; citation_volume=29; citation_publication_date=2009; citation_pages=8335-8349; citation_id=CR121"/> <meta name="citation_reference" content="Vargas JP, Lopez JC, Portavella M (2009) What are the functions of fish brain pallium? Brain Res Bull 79:436–440"/> <meta name="citation_reference" content="citation_journal_title=Nat Commun; citation_title=Abstract rule neurons in the endbrain support intelligent behaviour in corvid songbirds; citation_author=L Veit, A Nieder; citation_volume=4; citation_publication_date=2013; citation_pages=2878; citation_id=CR122"/> <meta name="citation_reference" content="citation_title=Animal models of pain; citation_inbook_title=Wall and Melzack’s textbook of pain; citation_publication_date=2006; citation_id=CR123; citation_author=CJ Vierck; citation_publisher=Churchill"/> <meta name="citation_reference" content="citation_journal_title=Behav Brain Res; citation_title=Integration of auditory and visual information in human face discrimination in pigeons. Behavioral and anatomical study; citation_author=S Watanabe, S Masuda; citation_volume=207; citation_publication_date=2010; citation_pages=61-69; citation_id=CR124"/> <meta name="citation_reference" content="citation_journal_title=PLoS ONE; citation_title=Stress-induced antinociception in fish reversed by naloxone; citation_author=CPB Wolkers, BA Junior, L Menescal-de-Oliveira, A Hoffmann; citation_volume=8; citation_publication_date=2013; citation_pages=e71175; citation_id=CR125"/> <meta name="citation_reference" content="citation_journal_title=Neuroscience; citation_title=μ but not δ and κ opioid receptor involvement in ventrolateral orbital cortex opioid-evoked antinociception in formalin test rats; citation_author=YF Xie, J Wang, FQ Huo, H Jia, JS Tang; citation_volume=126; citation_publication_date=2004; citation_pages=717-726; citation_id=CR126"/> <meta name="citation_reference" content="citation_journal_title=Brain Res; citation_title=Visual function in goldfish with unilateral and bilateral tectal ablation; citation_author=D Yager, SC Sharma, BG Grover; citation_volume=137; citation_publication_date=1977; citation_pages=267-275; citation_id=CR127"/> <meta name="citation_reference" content="citation_journal_title=Science; citation_title=Analgesia mediated by a direct spinal action of narcotics; citation_author=TL Yaksh, TA Rudy; citation_volume=192; citation_publication_date=1976; citation_pages=1357-1358; citation_id=CR128"/> <meta name="citation_reference" content="citation_journal_title=J Comp Neurol; citation_title=Visual, lateral line, and auditory ascending pathways to the dorsal telencephalic area through the rostrolateral region of the lateral preglomerular nucleus in Cyprinids; citation_author=N Yamamoto, H Ito; citation_volume=508; citation_publication_date=2008; citation_pages=615-647; citation_id=CR129"/> <meta name="citation_author" content="Key, Brian"/> <meta name="citation_author_email" content="brian.key@uq.edu.au"/> <meta name="citation_author_institution" content="School of Biomedical Sciences, University of Queensland, Brisbane, Australia"/> <meta name="format-detection" content="telephone=no"/> <meta name="citation_cover_date" content="2015/03/01"/> <meta property="og:url" content="https://link.springer.com/article/10.1007/s10539-014-9469-4"/> <meta property="og:type" content="article"/> <meta property="og:site_name" content="SpringerLink"/> <meta property="og:title" content="Fish do not feel pain and its implications for understanding phenomenal consciousness - Biology & Philosophy"/> <meta property="og:description" content="Phenomenal consciousness or the subjective experience of feeling sensory stimuli is fundamental to human existence. Because of the ubiquity of their subjective experiences, humans seem to readily accept the anthropomorphic extension of these mental states to other animals. Humans will typically extrapolate feelings of pain to animals if they respond physiologically and behaviourally to noxious stimuli. The alternative view that fish instead respond to noxious stimuli reflexly and with a limited behavioural repertoire is defended within the context of our current understanding of the neuroanatomy and neurophysiology of mental states. Consequently, a set of fundamental properties of neural tissue necessary for feeling pain or experiencing affective states in vertebrates is proposed. While mammals and birds possess the prerequisite neural architecture for phenomenal consciousness, it is concluded that fish lack these essential characteristics and hence do not feel pain."/> <meta property="og:image" content="https://media.springernature.com/full/springer-static/cover-hires/journal/10539"/> <meta name="format-detection" content="telephone=no"> <link rel="apple-touch-icon" sizes="180x180" href=/oscar-static/img/favicons/darwin/apple-touch-icon-92e819bf8a.png> <link rel="icon" type="image/png" sizes="192x192" href=/oscar-static/img/favicons/darwin/android-chrome-192x192-6f081ca7e5.png> <link rel="icon" type="image/png" sizes="32x32" href=/oscar-static/img/favicons/darwin/favicon-32x32-1435da3e82.png> <link rel="icon" type="image/png" sizes="16x16" href=/oscar-static/img/favicons/darwin/favicon-16x16-ed57f42bd2.png> <link rel="shortcut icon" data-test="shortcut-icon" href=/oscar-static/img/favicons/darwin/favicon-c6d59aafac.ico> <meta name="theme-color" content="#e6e6e6">   <link rel="stylesheet" media="print" href=/oscar-static/app-springerlink/css/print-b8af42253b.css> <style> html{text-size-adjust:100%;line-height:1.15}body{font-family:Merriweather Sans,Helvetica Neue,Helvetica,Arial,sans-serif;line-height:1.8;margin:0}details,main{display:block}h1{font-size:2em;margin:.67em 0}a{background-color:transparent;color:#025e8d}sub{bottom:-.25em;font-size:75%;line-height:0;position:relative;vertical-align:baseline}img{border:0;height:auto;max-width:100%;vertical-align:middle}button,input{font-family:inherit;font-size:100%;line-height:1.15;margin:0;overflow:visible}button{text-transform:none}[type=button],[type=submit],button{-webkit-appearance:button}[type=search]{-webkit-appearance:textfield;outline-offset:-2px}summary{display:list-item}[hidden]{display:none}button{cursor:pointer}svg{height:1rem;width:1rem} </style> <style>@media only print, only all and (prefers-color-scheme: no-preference), only all and (prefers-color-scheme: light), only all and (prefers-color-scheme: dark) { body{background:#fff;color:#222;font-family:Merriweather Sans,Helvetica Neue,Helvetica,Arial,sans-serif;line-height:1.8;min-height:100%}a{color:#025e8d;text-decoration:underline;text-decoration-skip-ink:auto}button{cursor:pointer}img{border:0;height:auto;max-width:100%;vertical-align:middle}html{box-sizing:border-box;font-size:100%;height:100%;overflow-y:scroll}h1{font-size:2.25rem}h2{font-size:1.75rem}h1,h2,h4{font-weight:700;line-height:1.2}h4{font-size:1.25rem}body{font-size:1.125rem}*{box-sizing:inherit}p{margin-bottom:2rem;margin-top:0}p:last-of-type{margin-bottom:0}.c-ad{text-align:center}@media only screen and (min-width:480px){.c-ad{padding:8px}}.c-ad--728x90{display:none}.c-ad--728x90 .c-ad__inner{min-height:calc(1.5em + 94px)}@media only screen and (min-width:876px){.js .c-ad--728x90{display:none}}.c-ad__label{color:#333;font-size:.875rem;font-weight:400;line-height:1.5;margin-bottom:4px}.c-ad__label,.c-status-message{font-family:Merriweather Sans,Helvetica Neue,Helvetica,Arial,sans-serif}.c-status-message{align-items:center;box-sizing:border-box;display:flex;position:relative;width:100%}.c-status-message :last-child{margin-bottom:0}.c-status-message--boxed{background-color:#fff;border:1px solid #ccc;line-height:1.4;padding:16px}.c-status-message__heading{font-family:Merriweather Sans,Helvetica Neue,Helvetica,Arial,sans-serif;font-size:.875rem;font-weight:700}.c-status-message__icon{fill:currentcolor;display:inline-block;flex:0 0 auto;height:1.5em;margin-right:8px;transform:translate(0);vertical-align:text-top;width:1.5em}.c-status-message__icon--top{align-self:flex-start}.c-status-message--info .c-status-message__icon{color:#003f8d}.c-status-message--boxed.c-status-message--info{border-bottom:4px solid #003f8d}.c-status-message--error .c-status-message__icon{color:#c40606}.c-status-message--boxed.c-status-message--error{border-bottom:4px solid #c40606}.c-status-message--success .c-status-message__icon{color:#00b8b0}.c-status-message--boxed.c-status-message--success{border-bottom:4px solid #00b8b0}.c-status-message--warning .c-status-message__icon{color:#edbc53}.c-status-message--boxed.c-status-message--warning{border-bottom:4px solid #edbc53}.eds-c-header{background-color:#fff;border-bottom:2px solid #01324b;font-family:Merriweather Sans,Helvetica Neue,Helvetica,Arial,sans-serif;font-size:1rem;line-height:1.5;padding:8px 0 0}.eds-c-header__container{align-items:center;display:flex;flex-wrap:nowrap;gap:8px 16px;justify-content:space-between;margin:0 auto 8px;max-width:1280px;padding:0 8px;position:relative}.eds-c-header__nav{border-top:2px solid #c5e0f4;padding-top:4px;position:relative}.eds-c-header__nav-container{align-items:center;display:flex;flex-wrap:wrap;margin:0 auto 4px;max-width:1280px;padding:0 8px;position:relative}.eds-c-header__nav-container>:not(:last-child){margin-right:32px}.eds-c-header__link-container{align-items:center;display:flex;flex:1 0 auto;gap:8px 16px;justify-content:space-between}.eds-c-header__list{list-style:none;margin:0;padding:0}.eds-c-header__list-item{font-weight:700;margin:0 auto;max-width:1280px;padding:8px}.eds-c-header__list-item:not(:last-child){border-bottom:2px solid #c5e0f4}.eds-c-header__item{color:inherit}@media only screen and (min-width:768px){.eds-c-header__item--menu{display:none;visibility:hidden}.eds-c-header__item--menu:first-child+*{margin-block-start:0}}.eds-c-header__item--inline-links{display:none;visibility:hidden}@media only screen and (min-width:768px){.eds-c-header__item--inline-links{display:flex;gap:16px 16px;visibility:visible}}.eds-c-header__item--divider:before{border-left:2px solid #c5e0f4;content:"";height:calc(100% - 16px);margin-left:-15px;position:absolute;top:8px}.eds-c-header__brand{padding:16px 8px}.eds-c-header__brand a{display:block;line-height:1;text-decoration:none}.eds-c-header__brand img{height:1.5rem;width:auto}.eds-c-header__link{color:inherit;display:inline-block;font-weight:700;padding:16px 8px;position:relative;text-decoration-color:transparent;white-space:nowrap;word-break:normal}.eds-c-header__icon{fill:currentcolor;display:inline-block;font-size:1.5rem;height:1em;transform:translate(0);vertical-align:bottom;width:1em}.eds-c-header__icon+*{margin-left:8px}.eds-c-header__expander{background-color:#f0f7fc}.eds-c-header__search{display:block;padding:24px 0}@media only screen and (min-width:768px){.eds-c-header__search{max-width:70%}}.eds-c-header__search-container{position:relative}.eds-c-header__search-label{color:inherit;display:inline-block;font-weight:700;margin-bottom:8px}.eds-c-header__search-input{background-color:#fff;border:1px solid #000;padding:8px 48px 8px 8px;width:100%}.eds-c-header__search-button{background-color:transparent;border:0;color:inherit;height:100%;padding:0 8px;position:absolute;right:0}.has-tethered.eds-c-header__expander{border-bottom:2px solid #01324b;left:0;margin-top:-2px;top:100%;width:100%;z-index:10}@media only screen and (min-width:768px){.has-tethered.eds-c-header__expander--menu{display:none;visibility:hidden}}.has-tethered .eds-c-header__heading{display:none;visibility:hidden}.has-tethered .eds-c-header__heading:first-child+*{margin-block-start:0}.has-tethered .eds-c-header__search{margin:auto}.eds-c-header__heading{margin:0 auto;max-width:1280px;padding:16px 16px 0}.eds-c-pagination{align-items:center;display:flex;flex-wrap:wrap;font-family:Merriweather Sans,Helvetica Neue,Helvetica,Arial,sans-serif;font-size:.875rem;gap:16px 0;justify-content:center;line-height:1.4;list-style:none;margin:0;padding:32px 0}@media only screen and (min-width:480px){.eds-c-pagination{padding:32px 16px}}.eds-c-pagination__item{margin-right:8px}.eds-c-pagination__item--prev{margin-right:16px}.eds-c-pagination__item--next .eds-c-pagination__link,.eds-c-pagination__item--prev .eds-c-pagination__link{padding:16px 8px}.eds-c-pagination__item--next{margin-left:8px}.eds-c-pagination__item:last-child{margin-right:0}.eds-c-pagination__link{align-items:center;color:#222;cursor:pointer;display:inline-block;font-size:1rem;margin:0;padding:16px 24px;position:relative;text-align:center;transition:all .2s ease 0s}.eds-c-pagination__link:visited{color:#222}.eds-c-pagination__link--disabled{border-color:#555;color:#555;cursor:default}.eds-c-pagination__link--active{background-color:#01324b;background-image:none;border-radius:8px;color:#fff}.eds-c-pagination__link--active:focus,.eds-c-pagination__link--active:hover,.eds-c-pagination__link--active:visited{color:#fff}.eds-c-pagination__link-container{align-items:center;display:flex}.eds-c-pagination__icon{fill:#222;height:1.5rem;width:1.5rem}.eds-c-pagination__icon--disabled{fill:#555}.eds-c-pagination__visually-hidden{clip:rect(0,0,0,0);border:0;clip-path:inset(50%);height:1px;overflow:hidden;padding:0;position:absolute!important;white-space:nowrap;width:1px}.c-breadcrumbs{color:#333;font-family:Merriweather Sans,Helvetica Neue,Helvetica,Arial,sans-serif;font-size:1rem;list-style:none;margin:0;padding:0}.c-breadcrumbs>li{display:inline}svg.c-breadcrumbs__chevron{fill:#333;height:10px;margin:0 .25rem;width:10px}.c-breadcrumbs--contrast,.c-breadcrumbs--contrast .c-breadcrumbs__link{color:#fff}.c-breadcrumbs--contrast svg.c-breadcrumbs__chevron{fill:#fff}@media only screen and (max-width:479px){.c-breadcrumbs .c-breadcrumbs__item{display:none}.c-breadcrumbs .c-breadcrumbs__item:last-child,.c-breadcrumbs .c-breadcrumbs__item:nth-last-child(2){display:inline}}.c-skip-link{background:#01324b;bottom:auto;color:#fff;font-family:Merriweather Sans,Helvetica Neue,Helvetica,Arial,sans-serif;font-size:1rem;padding:8px;position:absolute;text-align:center;transform:translateY(-100%);width:100%;z-index:9999}@media (prefers-reduced-motion:reduce){.c-skip-link{transition:top .3s ease-in-out 0s}}@media print{.c-skip-link{display:none}}.c-skip-link:active,.c-skip-link:hover,.c-skip-link:link,.c-skip-link:visited{color:#fff}.c-skip-link:focus{transform:translateY(0)}.l-with-sidebar{display:flex;flex-wrap:wrap}.l-with-sidebar>*{margin:0}.l-with-sidebar__sidebar{flex-basis:var(--with-sidebar--basis,400px);flex-grow:1}.l-with-sidebar>:not(.l-with-sidebar__sidebar){flex-basis:0px;flex-grow:999;min-width:var(--with-sidebar--min,53%)}.l-with-sidebar>:first-child{padding-right:4rem}@supports (gap:1em){.l-with-sidebar>:first-child{padding-right:0}.l-with-sidebar{gap:var(--with-sidebar--gap,4rem)}}.c-header__link{color:inherit;display:inline-block;font-weight:700;padding:16px 8px;position:relative;text-decoration-color:transparent;white-space:nowrap;word-break:normal}.app-masthead__colour-4{--background-color:#ff9500;--gradient-light:rgba(0,0,0,.5);--gradient-dark:rgba(0,0,0,.8)}.app-masthead{background:var(--background-color,#0070a8);position:relative}.app-masthead:after{background:radial-gradient(circle at top right,var(--gradient-light,rgba(0,0,0,.4)),var(--gradient-dark,rgba(0,0,0,.7)));bottom:0;content:"";left:0;position:absolute;right:0;top:0}@media only screen and (max-width:479px){.app-masthead:after{background:linear-gradient(225deg,var(--gradient-light,rgba(0,0,0,.4)),var(--gradient-dark,rgba(0,0,0,.7)))}}.app-masthead__container{color:var(--masthead-color,#fff);margin:0 auto;max-width:1280px;padding:0 16px;position:relative;z-index:1}.u-button{align-items:center;background-color:#01324b;background-image:none;border:4px solid transparent;border-radius:32px;cursor:pointer;display:inline-flex;font-family:Merriweather Sans,Helvetica Neue,Helvetica,Arial,sans-serif;font-size:.875rem;font-weight:700;justify-content:center;line-height:1.3;margin:0;padding:16px 32px;position:relative;transition:all .2s ease 0s;width:auto}.u-button svg,.u-button--contrast svg,.u-button--primary svg,.u-button--secondary svg,.u-button--tertiary svg{fill:currentcolor}.u-button,.u-button:visited{color:#fff}.u-button,.u-button:hover{box-shadow:0 0 0 1px #01324b;text-decoration:none}.u-button:hover{border:4px solid #fff}.u-button:focus{border:4px solid #fc0;box-shadow:none;outline:0;text-decoration:none}.u-button:focus,.u-button:hover{background-color:#fff;background-image:none;color:#01324b}.app-masthead--pastel .c-pdf-download .u-button--primary:focus svg path,.app-masthead--pastel .c-pdf-download .u-button--primary:hover svg path,.c-context-bar--sticky .c-context-bar__container .c-pdf-download .u-button--primary:focus svg path,.c-context-bar--sticky .c-context-bar__container .c-pdf-download .u-button--primary:hover svg path,.u-button--primary:focus svg path,.u-button--primary:hover svg path,.u-button:focus svg path,.u-button:hover svg path{fill:#01324b}.u-button--primary{background-color:#01324b;background-image:none;border:4px solid transparent;box-shadow:0 0 0 1px #01324b;color:#fff;font-weight:700}.u-button--primary:visited{color:#fff}.u-button--primary:hover{border:4px solid #fff;box-shadow:0 0 0 1px #01324b;text-decoration:none}.u-button--primary:focus{border:4px solid #fc0;box-shadow:none;outline:0;text-decoration:none}.u-button--primary:focus,.u-button--primary:hover{background-color:#fff;background-image:none;color:#01324b}.u-button--secondary{background-color:#fff;border:4px solid #fff;color:#01324b;font-weight:700}.u-button--secondary:visited{color:#01324b}.u-button--secondary:hover{border:4px solid #01324b;box-shadow:none}.u-button--secondary:focus,.u-button--secondary:hover{background-color:#01324b;color:#fff}.app-masthead--pastel .c-pdf-download .u-button--secondary:focus svg path,.app-masthead--pastel .c-pdf-download .u-button--secondary:hover svg path,.c-context-bar--sticky .c-context-bar__container .c-pdf-download .u-button--secondary:focus svg path,.c-context-bar--sticky .c-context-bar__container .c-pdf-download .u-button--secondary:hover svg path,.u-button--secondary:focus svg path,.u-button--secondary:hover svg path,.u-button--tertiary:focus svg path,.u-button--tertiary:hover svg path{fill:#fff}.u-button--tertiary{background-color:#ebf1f5;border:4px solid transparent;box-shadow:none;color:#666;font-weight:700}.u-button--tertiary:visited{color:#666}.u-button--tertiary:hover{border:4px solid #01324b;box-shadow:none}.u-button--tertiary:focus,.u-button--tertiary:hover{background-color:#01324b;color:#fff}.u-button--contrast{background-color:transparent;background-image:none;color:#fff;font-weight:400}.u-button--contrast:visited{color:#fff}.u-button--contrast,.u-button--contrast:focus,.u-button--contrast:hover{border:4px solid #fff}.u-button--contrast:focus,.u-button--contrast:hover{background-color:#fff;background-image:none;color:#000}.u-button--contrast:focus svg path,.u-button--contrast:hover svg path{fill:#000}.u-button--disabled,.u-button:disabled{background-color:transparent;background-image:none;border:4px solid #ccc;color:#000;cursor:default;font-weight:400;opacity:.7}.u-button--disabled svg,.u-button:disabled svg{fill:currentcolor}.u-button--disabled:visited,.u-button:disabled:visited{color:#000}.u-button--disabled:focus,.u-button--disabled:hover,.u-button:disabled:focus,.u-button:disabled:hover{border:4px solid #ccc;text-decoration:none}.u-button--disabled:focus,.u-button--disabled:hover,.u-button:disabled:focus,.u-button:disabled:hover{background-color:transparent;background-image:none;color:#000}.u-button--disabled:focus svg path,.u-button--disabled:hover svg path,.u-button:disabled:focus svg path,.u-button:disabled:hover svg path{fill:#000}.u-button--small,.u-button--xsmall{font-size:.875rem;padding:2px 8px}.u-button--small{padding:8px 16px}.u-button--large{font-size:1.125rem;padding:10px 35px}.u-button--full-width{display:flex;width:100%}.u-button--icon-left svg{margin-right:8px}.u-button--icon-right svg{margin-left:8px}.u-clear-both{clear:both}.u-container{margin:0 auto;max-width:1280px;padding:0 16px}.u-justify-content-space-between{justify-content:space-between}.u-display-none{display:none}.js .u-js-hide,.u-hide{display:none;visibility:hidden}.u-visually-hidden{clip:rect(0,0,0,0);border:0;clip-path:inset(50%);height:1px;overflow:hidden;padding:0;position:absolute!important;white-space:nowrap;width:1px}.u-icon{fill:currentcolor;display:inline-block;height:1em;transform:translate(0);vertical-align:text-top;width:1em}.u-list-reset{list-style:none;margin:0;padding:0}.u-ma-16{margin:16px}.u-mt-0{margin-top:0}.u-mt-24{margin-top:24px}.u-mt-32{margin-top:32px}.u-mb-8{margin-bottom:8px}.u-mb-32{margin-bottom:32px}.u-button-reset{background-color:transparent;border:0;padding:0}.u-sans-serif{font-family:Merriweather Sans,Helvetica Neue,Helvetica,Arial,sans-serif}.u-serif{font-family:Merriweather,serif}h1,h2,h4{-webkit-font-smoothing:antialiased}p{overflow-wrap:break-word;word-break:break-word}.u-h4{font-size:1.25rem;font-weight:700;line-height:1.2}.u-mbs-0{margin-block-start:0!important}.c-article-header{font-family:Merriweather Sans,Helvetica Neue,Helvetica,Arial,sans-serif}.c-article-identifiers{color:#6f6f6f;display:flex;flex-wrap:wrap;font-size:1rem;line-height:1.3;list-style:none;margin:0 0 8px;padding:0}.c-article-identifiers__item{border-right:1px solid #6f6f6f;list-style:none;margin-right:8px;padding-right:8px}.c-article-identifiers__item:last-child{border-right:0;margin-right:0;padding-right:0}@media only screen and (min-width:876px){.c-article-title{font-size:1.875rem;line-height:1.2}}.c-article-author-list{display:inline;font-size:1rem;list-style:none;margin:0 8px 0 0;padding:0;width:100%}.c-article-author-list__item{display:inline;padding-right:0}.c-article-author-list__show-more{display:none;margin-right:4px}.c-article-author-list__button,.js .c-article-author-list__item--hide,.js .c-article-author-list__show-more{display:none}.js .c-article-author-list--long .c-article-author-list__show-more,.js .c-article-author-list--long+.c-article-author-list__button{display:inline}@media only screen and (max-width:767px){.js .c-article-author-list__item--hide-small-screen{display:none}.js .c-article-author-list--short .c-article-author-list__show-more,.js .c-article-author-list--short+.c-article-author-list__button{display:inline}}#uptodate-client,.js .c-article-author-list--expanded .c-article-author-list__show-more{display:none!important}.js .c-article-author-list--expanded .c-article-author-list__item--hide-small-screen{display:inline!important}.c-article-author-list__button,.c-button-author-list{background:#ebf1f5;border:4px solid #ebf1f5;border-radius:20px;color:#666;font-size:.875rem;line-height:1.4;padding:2px 11px 2px 8px;text-decoration:none}.c-article-author-list__button svg,.c-button-author-list svg{margin:1px 4px 0 0}.c-article-author-list__button:hover,.c-button-author-list:hover{background:#025e8d;border-color:transparent;color:#fff}.c-article-body .c-article-access-provider{padding:8px 16px}.c-article-body .c-article-access-provider,.c-notes{border:1px solid #d5d5d5;border-image:initial;border-left:none;border-right:none;margin:24px 0}.c-article-body .c-article-access-provider__text{color:#555}.c-article-body .c-article-access-provider__text,.c-notes__text{font-size:1rem;margin-bottom:0;padding-bottom:2px;padding-top:2px;text-align:center}.c-article-body .c-article-author-affiliation__address{color:inherit;font-weight:700;margin:0}.c-article-body .c-article-author-affiliation__authors-list{list-style:none;margin:0;padding:0}.c-article-body .c-article-author-affiliation__authors-item{display:inline;margin-left:0}.c-article-authors-search{margin-bottom:24px;margin-top:0}.c-article-authors-search__item,.c-article-authors-search__title{font-family:Merriweather Sans,Helvetica Neue,Helvetica,Arial,sans-serif}.c-article-authors-search__title{color:#626262;font-size:1.05rem;font-weight:700;margin:0;padding:0}.c-article-authors-search__item{font-size:1rem}.c-article-authors-search__text{margin:0}.c-code-block{border:1px solid #fff;font-family:monospace;margin:0 0 24px;padding:20px}.c-code-block__heading{font-weight:400;margin-bottom:16px}.c-code-block__line{display:block;overflow-wrap:break-word;white-space:pre-wrap}.c-article-share-box{font-family:Merriweather Sans,Helvetica Neue,Helvetica,Arial,sans-serif;margin-bottom:24px}.c-article-share-box__description{font-size:1rem;margin-bottom:8px}.c-article-share-box__no-sharelink-info{font-size:.813rem;font-weight:700;margin-bottom:24px;padding-top:4px}.c-article-share-box__only-read-input{border:1px solid #d5d5d5;box-sizing:content-box;display:inline-block;font-size:.875rem;font-weight:700;height:24px;margin-bottom:8px;padding:8px 10px}.c-article-share-box__additional-info{color:#626262;font-size:.813rem}.c-article-share-box__button{background:#fff;box-sizing:content-box;text-align:center}.c-article-share-box__button--link-like{background-color:transparent;border:0;color:#025e8d;cursor:pointer;font-size:.875rem;margin-bottom:8px;margin-left:10px}.c-article-associated-content__container .c-article-associated-content__collection-label{font-size:.875rem;line-height:1.4}.c-article-associated-content__container .c-article-associated-content__collection-title{line-height:1.3}.c-reading-companion{clear:both;min-height:389px}.c-reading-companion__figures-list,.c-reading-companion__references-list{list-style:none;min-height:389px;padding:0}.c-reading-companion__references-list--numeric{list-style:decimal inside}.c-reading-companion__figure-item{border-top:1px solid #d5d5d5;font-size:1rem;padding:16px 8px 16px 0}.c-reading-companion__figure-item:first-child{border-top:none;padding-top:8px}.c-reading-companion__reference-item{font-size:1rem}.c-reading-companion__reference-item:first-child{border-top:none}.c-reading-companion__reference-item a{word-break:break-word}.c-reading-companion__reference-citation{display:inline}.c-reading-companion__reference-links{font-size:.813rem;font-weight:700;list-style:none;margin:8px 0 0;padding:0;text-align:right}.c-reading-companion__reference-links>a{display:inline-block;padding-left:8px}.c-reading-companion__reference-links>a:first-child{display:inline-block;padding-left:0}.c-reading-companion__figure-title{display:block;font-size:1.25rem;font-weight:700;line-height:1.2;margin:0 0 8px}.c-reading-companion__figure-links{display:flex;justify-content:space-between;margin:8px 0 0}.c-reading-companion__figure-links>a{align-items:center;display:flex}.c-article-section__figure-caption{display:block;margin-bottom:8px;word-break:break-word}.c-article-section__figure .video,p.app-article-masthead__access--above-download{margin:0 0 16px}.c-article-section__figure-description{font-size:1rem}.c-article-section__figure-description>*{margin-bottom:0}.c-cod{display:block;font-size:1rem;width:100%}.c-cod__form{background:#ebf0f3}.c-cod__prompt{font-size:1.125rem;line-height:1.3;margin:0 0 24px}.c-cod__label{display:block;margin:0 0 4px}.c-cod__row{display:flex;margin:0 0 16px}.c-cod__row:last-child{margin:0}.c-cod__input{border:1px solid #d5d5d5;border-radius:2px;flex-shrink:0;margin:0;padding:13px}.c-cod__input--submit{background-color:#025e8d;border:1px solid #025e8d;color:#fff;flex-shrink:1;margin-left:8px;transition:background-color .2s ease-out 0s,color .2s ease-out 0s}.c-cod__input--submit-single{flex-basis:100%;flex-shrink:0;margin:0}.c-cod__input--submit:focus,.c-cod__input--submit:hover{background-color:#fff;color:#025e8d}.save-data .c-article-author-institutional-author__sub-division,.save-data .c-article-equation__number,.save-data .c-article-figure-description,.save-data .c-article-fullwidth-content,.save-data .c-article-main-column,.save-data .c-article-satellite-article-link,.save-data .c-article-satellite-subtitle,.save-data .c-article-table-container,.save-data .c-blockquote__body,.save-data .c-code-block__heading,.save-data .c-reading-companion__figure-title,.save-data .c-reading-companion__reference-citation,.save-data .c-site-messages--nature-briefing-email-variant .serif,.save-data .c-site-messages--nature-briefing-email-variant.serif,.save-data .serif,.save-data .u-serif,.save-data h1,.save-data h2,.save-data h3{font-family:Merriweather Sans,Helvetica Neue,Helvetica,Arial,sans-serif}.c-pdf-download__link{display:flex;flex:1 1 0%;padding:13px 24px}.c-pdf-download__link:hover{text-decoration:none}@media only screen and (min-width:768px){.c-context-bar--sticky .c-pdf-download__link{align-items:center;flex:1 1 183px}}@media only screen and (max-width:320px){.c-context-bar--sticky .c-pdf-download__link{padding:16px}}.c-article-body .c-article-recommendations-list,.c-book-body .c-article-recommendations-list{display:flex;flex-direction:row;gap:16px 16px;margin:0;max-width:100%;padding:16px 0 0}.c-article-body .c-article-recommendations-list__item,.c-book-body .c-article-recommendations-list__item{flex:1 1 0%}@media only screen and (max-width:767px){.c-article-body .c-article-recommendations-list,.c-book-body .c-article-recommendations-list{flex-direction:column}}.c-article-body .c-article-recommendations-card__authors{display:none;font-family:Merriweather Sans,Helvetica Neue,Helvetica,Arial,sans-serif;font-size:.875rem;line-height:1.5;margin:0 0 8px}@media only screen and (max-width:767px){.c-article-body .c-article-recommendations-card__authors{display:block;margin:0}}.c-article-body .c-article-history{margin-top:24px}.app-article-metrics-bar p{margin:0}.app-article-masthead{display:flex;flex-direction:column;gap:16px 16px;padding:16px 0 24px}.app-article-masthead__info{display:flex;flex-direction:column;flex-grow:1}.app-article-masthead__brand{border-top:1px solid hsla(0,0%,100%,.8);display:flex;flex-direction:column;flex-shrink:0;gap:8px 8px;min-height:96px;padding:16px 0 0}.app-article-masthead__brand img{border:1px solid #fff;border-radius:8px;box-shadow:0 4px 15px 0 hsla(0,0%,50%,.25);height:auto;left:0;position:absolute;width:72px}.app-article-masthead__journal-link{display:block;font-size:1.125rem;font-weight:700;margin:0 0 8px;max-width:400px;padding:0 0 0 88px;position:relative}.app-article-masthead__journal-title{-webkit-box-orient:vertical;-webkit-line-clamp:3;display:-webkit-box;overflow:hidden}.app-article-masthead__submission-link{align-items:center;display:flex;font-size:1rem;gap:4px 4px;margin:0 0 0 88px}.app-article-masthead__access{align-items:center;display:flex;flex-wrap:wrap;font-size:.875rem;font-weight:300;gap:4px 4px;margin:0}.app-article-masthead__buttons{display:flex;flex-flow:column wrap;gap:16px 16px}.app-article-masthead__access svg,.app-masthead--pastel .c-pdf-download .u-button--primary svg,.app-masthead--pastel .c-pdf-download .u-button--secondary svg,.c-context-bar--sticky .c-context-bar__container .c-pdf-download .u-button--primary svg,.c-context-bar--sticky .c-context-bar__container .c-pdf-download .u-button--secondary svg{fill:currentcolor}.app-article-masthead a{color:#fff}.app-masthead--pastel .c-pdf-download .u-button--primary,.c-context-bar--sticky .c-context-bar__container .c-pdf-download .u-button--primary{background-color:#025e8d;background-image:none;border:2px solid transparent;box-shadow:none;color:#fff;font-weight:700}.app-masthead--pastel .c-pdf-download .u-button--primary:visited,.c-context-bar--sticky .c-context-bar__container .c-pdf-download .u-button--primary:visited{color:#fff}.app-masthead--pastel .c-pdf-download .u-button--primary:hover,.c-context-bar--sticky .c-context-bar__container .c-pdf-download .u-button--primary:hover{text-decoration:none}.app-masthead--pastel .c-pdf-download .u-button--primary:focus,.c-context-bar--sticky .c-context-bar__container .c-pdf-download .u-button--primary:focus{border:4px solid #fc0;box-shadow:none;outline:0;text-decoration:none}.app-masthead--pastel .c-pdf-download .u-button--primary:focus,.app-masthead--pastel .c-pdf-download .u-button--primary:hover,.c-context-bar--sticky .c-context-bar__container .c-pdf-download .u-button--primary:focus,.c-context-bar--sticky .c-context-bar__container .c-pdf-download .u-button--primary:hover{background-color:#fff;background-image:none;color:#01324b}.app-masthead--pastel .c-pdf-download .u-button--primary:hover,.c-context-bar--sticky .c-context-bar__container .c-pdf-download .u-button--primary:hover{background:0 0;border:2px solid #025e8d;box-shadow:none;color:#025e8d}.app-masthead--pastel .c-pdf-download .u-button--secondary,.c-context-bar--sticky .c-context-bar__container .c-pdf-download .u-button--secondary{background:0 0;border:2px solid #025e8d;color:#025e8d;font-weight:700}.app-masthead--pastel .c-pdf-download .u-button--secondary:visited,.c-context-bar--sticky .c-context-bar__container .c-pdf-download .u-button--secondary:visited{color:#01324b}.app-masthead--pastel .c-pdf-download .u-button--secondary:hover,.c-context-bar--sticky .c-context-bar__container .c-pdf-download .u-button--secondary:hover{background-color:#01324b;background-color:#025e8d;border:2px solid transparent;box-shadow:none;color:#fff}.app-masthead--pastel .c-pdf-download .u-button--secondary:focus,.c-context-bar--sticky .c-context-bar__container .c-pdf-download .u-button--secondary:focus{background-color:#fff;background-image:none;border:4px solid #fc0;color:#01324b}@media only screen and (min-width:768px){.app-article-masthead{flex-direction:row;gap:64px 64px;padding:24px 0}.app-article-masthead__brand{border:0;padding:0}.app-article-masthead__brand img{height:auto;position:static;width:auto}.app-article-masthead__buttons{align-items:center;flex-direction:row;margin-top:auto}.app-article-masthead__journal-link{display:flex;flex-direction:column;gap:24px 24px;margin:0 0 8px;padding:0}.app-article-masthead__submission-link{margin:0}}@media only screen and (min-width:1024px){.app-article-masthead__brand{flex-basis:400px}}.app-article-masthead .c-article-identifiers{font-size:.875rem;font-weight:300;line-height:1;margin:0 0 8px;overflow:hidden;padding:0}.app-article-masthead .c-article-identifiers--cite-list{margin:0 0 16px}.app-article-masthead .c-article-identifiers *{color:#fff}.app-article-masthead .c-cod{display:none}.app-article-masthead .c-article-identifiers__item{border-left:1px solid #fff;border-right:0;margin:0 17px 8px -9px;padding:0 0 0 8px}.app-article-masthead .c-article-identifiers__item--cite{border-left:0}.app-article-metrics-bar{display:flex;flex-wrap:wrap;font-size:1rem;padding:16px 0 0;row-gap:24px}.app-article-metrics-bar__item{padding:0 16px 0 0}.app-article-metrics-bar__count{font-weight:700}.app-article-metrics-bar__label{font-weight:400;padding-left:4px}.app-article-metrics-bar__icon{height:auto;margin-right:4px;margin-top:-4px;width:auto}.app-article-metrics-bar__arrow-icon{margin:4px 0 0 4px}.app-article-metrics-bar a{color:#000}.app-article-metrics-bar .app-article-metrics-bar__item--metrics{padding-right:0}.app-overview-section .c-article-author-list,.app-overview-section__authors{line-height:2}.app-article-metrics-bar{margin-top:8px}.c-book-toc-pagination+.c-book-section__back-to-top{margin-top:0}.c-article-body .c-article-access-provider__text--chapter{color:#222;font-family:Merriweather Sans,Helvetica Neue,Helvetica,Arial,sans-serif;padding:20px 0}.c-article-body .c-article-access-provider__text--chapter svg.c-status-message__icon{fill:#003f8d;vertical-align:middle}.c-article-body-section__content--separator{padding-top:40px}.c-pdf-download__link{max-height:44px}.app-article-access .u-button--primary,.app-article-access .u-button--primary:visited{color:#fff}.c-article-sidebar{display:none}@media only screen and (min-width:1024px){.c-article-sidebar{display:block}}.c-cod__form{border-radius:12px}.c-cod__label{font-size:.875rem}.c-cod .c-status-message{align-items:center;justify-content:center;margin-bottom:16px;padding-bottom:16px}@media only screen and (min-width:1024px){.c-cod .c-status-message{align-items:inherit}}.c-cod .c-status-message__icon{margin-top:4px}.c-cod .c-cod__prompt{font-size:1rem;margin-bottom:16px}.c-article-body .app-article-access,.c-book-body .app-article-access{display:block}@media only screen and (min-width:1024px){.c-article-body .app-article-access,.c-book-body .app-article-access{display:none}}.c-article-body .app-card-service{margin-bottom:32px}@media only screen and (min-width:1024px){.c-article-body .app-card-service{display:none}}.app-article-access .buybox__buy .u-button--secondary,.app-article-access .u-button--primary,.c-cod__row .u-button--primary{background-color:#025e8d;border:2px solid #025e8d;box-shadow:none;font-size:1rem;font-weight:700;gap:8px 8px;justify-content:center;line-height:1.5;padding:8px 24px}.app-article-access .buybox__buy .u-button--secondary,.app-article-access .u-button--primary:hover,.c-cod__row .u-button--primary:hover{background-color:#fff;color:#025e8d}.app-article-access .buybox__buy .u-button--secondary:hover{background-color:#025e8d;color:#fff}.buybox__buy .c-notes__text{color:#666;font-size:.875rem;padding:0 16px 8px}.c-cod__input{flex-basis:auto;width:100%}.c-article-title{font-family:Merriweather Sans,Helvetica Neue,Helvetica,Arial,sans-serif;font-size:2.25rem;font-weight:700;line-height:1.2;margin:12px 0}.c-reading-companion__figure-item figure{margin:0}@media only screen and (min-width:768px){.c-article-title{margin:16px 0}}.app-article-access{border:1px solid #c5e0f4;border-radius:12px}.app-article-access__heading{border-bottom:1px solid #c5e0f4;font-family:Merriweather Sans,Helvetica Neue,Helvetica,Arial,sans-serif;font-size:1.125rem;font-weight:700;margin:0;padding:16px;text-align:center}.app-article-access .buybox__info svg{vertical-align:middle}.c-article-body .app-article-access p{margin-bottom:0}.app-article-access .buybox__info{font-family:Merriweather Sans,Helvetica Neue,Helvetica,Arial,sans-serif;font-size:1rem;margin:0}.app-article-access{margin:0 0 32px}@media only screen and (min-width:1024px){.app-article-access{margin:0 0 24px}}.c-status-message{font-size:1rem}.c-article-body{font-size:1.125rem}.c-article-body dl,.c-article-body ol,.c-article-body p,.c-article-body ul{margin-bottom:32px;margin-top:0}.c-article-access-provider__text:last-of-type,.c-article-body .c-notes__text:last-of-type{margin-bottom:0}.c-article-body ol p,.c-article-body ul p{margin-bottom:16px}.c-article-section__figure-caption{font-family:Merriweather Sans,Helvetica Neue,Helvetica,Arial,sans-serif}.c-reading-companion__figure-item{border-top-color:#c5e0f4}.c-reading-companion__sticky{max-width:400px}.c-article-section .c-article-section__figure-description>*{font-size:1rem;margin-bottom:16px}.c-reading-companion__reference-item{border-top:1px solid #d5d5d5;padding:16px 0}.c-reading-companion__reference-item:first-child{padding-top:0}.c-article-share-box__button,.js .c-article-authors-search__item .c-article-button{background:0 0;border:2px solid #025e8d;border-radius:32px;box-shadow:none;color:#025e8d;font-size:1rem;font-weight:700;line-height:1.5;margin:0;padding:8px 24px;transition:all .2s ease 0s}.c-article-authors-search__item .c-article-button{width:100%}.c-pdf-download .u-button{background-color:#fff;border:2px solid #fff;color:#01324b;justify-content:center}.c-context-bar__container .c-pdf-download .u-button svg,.c-pdf-download .u-button svg{fill:currentcolor}.c-pdf-download .u-button:visited{color:#01324b}.c-pdf-download .u-button:hover{border:4px solid #01324b;box-shadow:none}.c-pdf-download .u-button:focus,.c-pdf-download .u-button:hover{background-color:#01324b}.c-pdf-download .u-button:focus svg path,.c-pdf-download .u-button:hover svg path{fill:#fff}.c-context-bar__container .c-pdf-download .u-button{background-image:none;border:2px solid;color:#fff}.c-context-bar__container .c-pdf-download .u-button:visited{color:#fff}.c-context-bar__container .c-pdf-download .u-button:hover{text-decoration:none}.c-context-bar__container .c-pdf-download .u-button:focus{box-shadow:none;outline:0;text-decoration:none}.c-context-bar__container .c-pdf-download .u-button:focus,.c-context-bar__container .c-pdf-download .u-button:hover{background-color:#fff;background-image:none;color:#01324b}.c-context-bar__container .c-pdf-download .u-button:focus svg path,.c-context-bar__container .c-pdf-download .u-button:hover svg path{fill:#01324b}.c-context-bar__container .c-pdf-download .u-button,.c-pdf-download .u-button{box-shadow:none;font-size:1rem;font-weight:700;line-height:1.5;padding:8px 24px}.c-context-bar__container .c-pdf-download .u-button{background-color:#025e8d}.c-pdf-download .u-button:hover{border:2px solid #fff}.c-pdf-download .u-button:focus,.c-pdf-download .u-button:hover{background:0 0;box-shadow:none;color:#fff}.c-context-bar__container .c-pdf-download .u-button:hover{border:2px solid #025e8d;box-shadow:none;color:#025e8d}.c-context-bar__container .c-pdf-download .u-button:focus,.c-pdf-download .u-button:focus{border:2px solid #025e8d}.c-article-share-box__button:focus:focus,.c-article__pill-button:focus:focus,.c-context-bar__container .c-pdf-download .u-button:focus:focus,.c-pdf-download .u-button:focus:focus{outline:3px solid #08c;will-change:transform}.c-pdf-download__link .u-icon{padding-top:0}.c-bibliographic-information__column button{margin-bottom:16px}.c-article-body .c-article-author-affiliation__list p,.c-article-body .c-article-author-information__list p,figure{margin:0}.c-article-share-box__button{margin-right:16px}.c-status-message--boxed{border-radius:12px}.c-article-associated-content__collection-title{font-size:1rem}.app-card-service__description,.c-article-body .app-card-service__description{color:#222;margin-bottom:0;margin-top:8px}.app-article-access__subscriptions a,.app-article-access__subscriptions a:visited,.app-book-series-listing__item a,.app-book-series-listing__item a:hover,.app-book-series-listing__item a:visited,.c-article-author-list a,.c-article-author-list a:visited,.c-article-buy-box a,.c-article-buy-box a:visited,.c-article-peer-review a,.c-article-peer-review a:visited,.c-article-satellite-subtitle a,.c-article-satellite-subtitle a:visited,.c-breadcrumbs__link,.c-breadcrumbs__link:hover,.c-breadcrumbs__link:visited{color:#000}.c-article-author-list svg{height:24px;margin:0 0 0 6px;width:24px}.c-article-header{margin-bottom:32px}@media only screen and (min-width:876px){.js .c-ad--conditional{display:block}}.u-lazy-ad-wrapper{background-color:#fff;display:none;min-height:149px}@media only screen and (min-width:876px){.u-lazy-ad-wrapper{display:block}}p.c-ad__label{margin-bottom:4px}.c-ad--728x90{background-color:#fff;border-bottom:2px solid #cedbe0} } </style> <style>@media only print, only all and (prefers-color-scheme: no-preference), only all and (prefers-color-scheme: light), only all and (prefers-color-scheme: dark) { .eds-c-header__brand img{height:24px;width:203px}.app-article-masthead__journal-link img{height:93px;width:72px}@media only screen and (min-width:769px){.app-article-masthead__journal-link img{height:161px;width:122px}} } </style> <link rel="stylesheet" data-test="critical-css-handler" data-inline-css-source="critical-css" href=/oscar-static/app-springerlink/css/core-darwin-5272567b64.css media="print" onload="this.media='all';this.onload=null"> <link rel="stylesheet" data-test="critical-css-handler" data-inline-css-source="critical-css" href="/oscar-static/app-springerlink/css/enhanced-darwin-article-72ba046d97.css" media="print" onload="this.media='only print, only all and (prefers-color-scheme: no-preference), only all and (prefers-color-scheme: light), only all and (prefers-color-scheme: dark)';this.onload=null"> <script type="text/javascript"> config = { env: 'live', site: '10539.springer.com', siteWithPath: '10539.springer.com' + window.location.pathname, twitterHashtag: '10539', cmsPrefix: 'https://studio-cms.springernature.com/studio/', publisherBrand: 'Springer', mustardcut: false }; </script> <script> window.dataLayer = [{"GA Key":"UA-26408784-1","DOI":"10.1007/s10539-014-9469-4","Page":"article","springerJournal":true,"Publishing Model":"Hybrid Access","page":{"attributes":{"environment":"live"}},"Country":"HK","japan":false,"doi":"10.1007-s10539-014-9469-4","Journal Id":10539,"Journal Title":"Biology \u0026 Philosophy","imprint":"Springer","Keywords":"Fish, Pain, Phenomenal consciousness, Affective states, Avoidance learning, Neocortex, Pallium","kwrd":["Fish","Pain","Phenomenal_consciousness","Affective_states","Avoidance_learning","Neocortex","Pallium"],"Labs":"Y","ksg":"Krux.segments","kuid":"Krux.uid","Has Body":"Y","Features":[],"Open Access":"Y","hasAccess":"Y","bypassPaywall":"N","user":{"license":{"businessPartnerID":[],"businessPartnerIDString":""}},"Access Type":"open","Bpids":"","Bpnames":"","BPID":["1"],"VG Wort Identifier":"vgzm.415900-10.1007-s10539-014-9469-4","Full HTML":"Y","Subject Codes":["SCE","SCE34010","SCL21001"],"pmc":["E","E34010","L21001"],"session":{"authentication":{"loginStatus":"N"},"attributes":{"edition":"academic"}},"content":{"serial":{"eissn":"1572-8404","pissn":"0169-3867"},"type":"Article","category":{"pmc":{"primarySubject":"Philosophy","primarySubjectCode":"E","secondarySubjects":{"1":"Philosophy of Biology","2":"Evolutionary Biology"},"secondarySubjectCodes":{"1":"E34010","2":"L21001"}},"sucode":"SC21","articleType":"Article"},"attributes":{"deliveryPlatform":"oscar"}},"Event Category":"Article"}]; </script> <script data-test="springer-link-article-datalayer"> window.dataLayer = window.dataLayer || []; window.dataLayer.push({ ga4MeasurementId: 'G-B3E4QL2TPR', ga360TrackingId: 'UA-26408784-1', twitterId: 'o47a7', baiduId: 'aef3043f025ccf2305af8a194652d70b', ga4ServerUrl: 'https://collect.springer.com', imprint: 'springerlink', page: { attributes:{ featureFlags: [{ name: 'darwin-orion', active: true }, { name: 'chapter-books-recs', active: true } ], darwinAvailable: true } } }); </script> <script> (function(w, d) { w.config = w.config || {}; w.config.mustardcut = false; if (w.matchMedia && w.matchMedia('only print, only all and (prefers-color-scheme: no-preference), only all and (prefers-color-scheme: light), only all and (prefers-color-scheme: dark)').matches) { w.config.mustardcut = true; d.classList.add('js'); d.classList.remove('grade-c'); d.classList.remove('no-js'); } })(window, document.documentElement); </script> <script class="js-entry"> if (window.config.mustardcut) { (function(w, d) { window.Component = {}; window.suppressShareButton = false; window.onArticlePage = true; var currentScript = d.currentScript || d.head.querySelector('script.js-entry'); function catchNoModuleSupport() { var scriptEl = d.createElement('script'); return (!('noModule' in scriptEl) && 'onbeforeload' in scriptEl) } var headScripts = [ {'src': '/oscar-static/js/polyfill-es5-bundle-572d4fec60.js', 'async': false} ]; var bodyScripts = [ {'src': '/oscar-static/js/global-article-es5-bundle-dad1690b0d.js', 'async': false, 'module': false}, {'src': '/oscar-static/js/global-article-es6-bundle-e7d03c4cb3.js', 'async': false, 'module': true} ]; function createScript(script) { var scriptEl = d.createElement('script'); scriptEl.src = script.src; scriptEl.async = script.async; if (script.module === true) { scriptEl.type = "module"; if (catchNoModuleSupport()) { scriptEl.src = ''; } } else if (script.module === false) { scriptEl.setAttribute('nomodule', true) } if (script.charset) { scriptEl.setAttribute('charset', script.charset); } return scriptEl; } for (var i = 0; i < headScripts.length; ++i) { var scriptEl = createScript(headScripts[i]); currentScript.parentNode.insertBefore(scriptEl, currentScript.nextSibling); } d.addEventListener('DOMContentLoaded', function() { for (var i = 0; i < bodyScripts.length; ++i) { var scriptEl = createScript(bodyScripts[i]); d.body.appendChild(scriptEl); } }); // Webfont repeat view var config = w.config; if (config && config.publisherBrand && sessionStorage.fontsLoaded === 'true') { d.documentElement.className += ' webfonts-loaded'; } })(window, document); } </script> <script data-src="https://cdn.optimizely.com/js/27195530232.js" data-cc-script="C03"></script> <script data-test="gtm-head"> window.initGTM = function() { if (window.config.mustardcut) { (function (w, d, s, l, i) { w[l] = w[l] || []; w[l].push({'gtm.start': new Date().getTime(), event: 'gtm.js'}); var f = d.getElementsByTagName(s)[0], j = d.createElement(s), dl = l != 'dataLayer' ? '&l=' + l : ''; j.async = true; j.src = 'https://www.googletagmanager.com/gtm.js?id=' + i + dl; f.parentNode.insertBefore(j, f); })(window, document, 'script', 'dataLayer', 'GTM-MRVXSHQ'); } } </script> <script> (function (w, d, t) { function cc() { var h = w.location.hostname; var e = d.createElement(t), s = d.getElementsByTagName(t)[0]; if (h.indexOf('springer.com') > -1 && h.indexOf('biomedcentral.com') === -1 && h.indexOf('springeropen.com') === -1) { if (h.indexOf('link-qa.springer.com') > -1 || h.indexOf('test-www.springer.com') > -1) { e.src = 'https://cmp.springer.com/production_live/en/consent-bundle-17-52.js'; e.setAttribute('onload', "initGTM(window,document,'script','dataLayer','GTM-MRVXSHQ')"); } else { e.src = 'https://cmp.springer.com/production_live/en/consent-bundle-17-52.js'; e.setAttribute('onload', "initGTM(window,document,'script','dataLayer','GTM-MRVXSHQ')"); } } else if (h.indexOf('biomedcentral.com') > -1) { if (h.indexOf('biomedcentral.com.qa') > -1) { e.src = 'https://cmp.biomedcentral.com/production_live/en/consent-bundle-15-36.js'; e.setAttribute('onload', "initGTM(window,document,'script','dataLayer','GTM-MRVXSHQ')"); } else { e.src = 'https://cmp.biomedcentral.com/production_live/en/consent-bundle-15-36.js'; e.setAttribute('onload', "initGTM(window,document,'script','dataLayer','GTM-MRVXSHQ')"); } } else if (h.indexOf('springeropen.com') > -1) { if (h.indexOf('springeropen.com.qa') > -1) { e.src = 'https://cmp.springernature.com/production_live/en/consent-bundle-16-34.js'; e.setAttribute('onload', "initGTM(window,document,'script','dataLayer','GTM-MRVXSHQ')"); } else { e.src = 'https://cmp.springernature.com/production_live/en/consent-bundle-16-34.js'; e.setAttribute('onload', "initGTM(window,document,'script','dataLayer','GTM-MRVXSHQ')"); } } else if (h.indexOf('springernature.com') > -1) { if (h.indexOf('beta-qa.springernature.com') > -1) { e.src = 'https://cmp.springernature.com/production_live/en/consent-bundle-49-43.js'; e.setAttribute('onload', "initGTM(window,document,'script','dataLayer','GTM-NK22KLS')"); } else { e.src = 'https://cmp.springernature.com/production_live/en/consent-bundle-49-43.js'; e.setAttribute('onload', "initGTM(window,document,'script','dataLayer','GTM-NK22KLS')"); } } else { e.src = '/oscar-static/js/cookie-consent-es5-bundle-cb57c2c98a.js'; e.setAttribute('data-consent', h); } s.insertAdjacentElement('afterend', e); } cc(); })(window, document, 'script'); </script> <link rel="canonical" href="https://link.springer.com/article/10.1007/s10539-014-9469-4"/> <script type="application/ld+json">{"mainEntity":{"headline":"Fish do not feel pain and its implications for understanding phenomenal consciousness","description":"Phenomenal consciousness or the subjective experience of feeling sensory stimuli is fundamental to human existence. Because of the ubiquity of their subjective experiences, humans seem to readily accept the anthropomorphic extension of these mental states to other animals. Humans will typically extrapolate feelings of pain to animals if they respond physiologically and behaviourally to noxious stimuli. The alternative view that fish instead respond to noxious stimuli reflexly and with a limited behavioural repertoire is defended within the context of our current understanding of the neuroanatomy and neurophysiology of mental states. Consequently, a set of fundamental properties of neural tissue necessary for feeling pain or experiencing affective states in vertebrates is proposed. While mammals and birds possess the prerequisite neural architecture for phenomenal consciousness, it is concluded that fish lack these essential characteristics and hence do not feel pain.","datePublished":"2014-12-16T00:00:00Z","dateModified":"2014-12-16T00:00:00Z","pageStart":"149","pageEnd":"165","sameAs":"https://doi.org/10.1007/s10539-014-9469-4","keywords":["Fish","Pain","Phenomenal consciousness","Affective states","Avoidance learning","Neocortex","Pallium","Philosophy of Biology","Evolutionary Biology"],"image":[],"isPartOf":{"name":"Biology & Philosophy","issn":["1572-8404","0169-3867"],"volumeNumber":"30","@type":["Periodical","PublicationVolume"]},"publisher":{"name":"Springer Netherlands","logo":{"url":"https://www.springernature.com/app-sn/public/images/logo-springernature.png","@type":"ImageObject"},"@type":"Organization"},"author":[{"name":"Brian Key","affiliation":[{"name":"University of Queensland","address":{"name":"School of Biomedical Sciences, University of Queensland, Brisbane, Australia","@type":"PostalAddress"},"@type":"Organization"}],"email":"brian.key@uq.edu.au","@type":"Person"}],"isAccessibleForFree":true,"@type":"ScholarlyArticle"},"@context":"https://schema.org","@type":"WebPage"}</script> </head> <body class="" >  <noscript> <iframe src="https://www.googletagmanager.com/ns.html?id=GTM-MRVXSHQ" height="0" width="0" style="display:none;visibility:hidden"></iframe> </noscript>   <noscript data-test="gtm-body"> <iframe src="https://www.googletagmanager.com/ns.html?id=GTM-MRVXSHQ" height="0" width="0" style="display:none;visibility:hidden"></iframe> </noscript>  <div class="u-visually-hidden" aria-hidden="true" data-test="darwin-icons"> <?xml version="1.0" encoding="UTF-8"?><!DOCTYPE svg PUBLIC "-//W3C//DTD SVG 1.1//EN" "http://www.w3.org/Graphics/SVG/1.1/DTD/svg11.dtd"><svg xmlns="http://www.w3.org/2000/svg" xmlns:xlink="http://www.w3.org/1999/xlink"><symbol id="icon-eds-i-accesses-medium" viewBox="0 0 24 24"><path d="M15.59 1a1 1 0 0 1 .706.291l5.41 5.385a1 1 0 0 1 .294.709v13.077c0 .674-.269 1.32-.747 1.796a2.549 2.549 0 0 1-1.798.742H15a1 1 0 0 1 0-2h4.455a.549.549 0 0 0 .387-.16.535.535 0 0 0 .158-.378V7.8L15.178 3H5.545a.543.543 0 0 0-.538.451L5 3.538v8.607a1 1 0 0 1-2 0V3.538A2.542 2.542 0 0 1 5.545 1h10.046ZM8 13c2.052 0 4.66 1.61 6.36 3.4l.124.141c.333.41.516.925.516 1.459 0 .6-.232 1.178-.64 1.599C12.666 21.388 10.054 23 8 23c-2.052 0-4.66-1.61-6.353-3.393A2.31 2.31 0 0 1 1 18c0-.6.232-1.178.64-1.6C3.34 14.61 5.948 13 8 13Zm0 2c-1.369 0-3.552 1.348-4.917 2.785A.31.31 0 0 0 3 18c0 .083.031.161.09.222C4.447 19.652 6.631 21 8 21c1.37 0 3.556-1.35 4.917-2.785A.31.31 0 0 0 13 18a.32.32 0 0 0-.048-.17l-.042-.052C11.553 16.348 9.369 15 8 15Zm0 1a2 2 0 1 1 0 4 2 2 0 0 1 0-4Z"/></symbol><symbol id="icon-eds-i-altmetric-medium" viewBox="0 0 24 24"><path d="M12 1c5.978 0 10.843 4.77 10.996 10.712l.004.306-.002.022-.002.248C22.843 18.23 17.978 23 12 23 5.925 23 1 18.075 1 12S5.925 1 12 1Zm-1.726 9.246L8.848 12.53a1 1 0 0 1-.718.461L8.003 13l-4.947.014a9.001 9.001 0 0 0 17.887-.001L16.553 13l-2.205 3.53a1 1 0 0 1-1.735-.068l-.05-.11-2.289-6.106ZM12 3a9.001 9.001 0 0 0-8.947 8.013l4.391-.012L9.652 7.47a1 1 0 0 1 1.784.179l2.288 6.104 1.428-2.283a1 1 0 0 1 .722-.462l.129-.008 4.943.012A9.001 9.001 0 0 0 12 3Z"/></symbol><symbol id="icon-eds-i-arrow-bend-down-medium" viewBox="0 0 24 24"><path d="m11.852 20.989.058.007L12 21l.075-.003.126-.017.111-.03.111-.044.098-.052.104-.074.082-.073 6-6a1 1 0 0 0-1.414-1.414L13 17.585v-12.2C13 4.075 11.964 3 10.667 3H4a1 1 0 1 0 0 2h6.667c.175 0 .333.164.333.385v12.2l-4.293-4.292a1 1 0 0 0-1.32-.083l-.094.083a1 1 0 0 0 0 1.414l6 6c.035.036.073.068.112.097l.11.071.114.054.105.035.118.025Z"/></symbol><symbol id="icon-eds-i-arrow-bend-down-small" viewBox="0 0 16 16"><path d="M1 2a1 1 0 0 0 1 1h5v8.585L3.707 8.293a1 1 0 0 0-1.32-.083l-.094.083a1 1 0 0 0 0 1.414l5 5 .063.059.093.069.081.048.105.048.104.035.105.022.096.01h.136l.122-.018.113-.03.103-.04.1-.053.102-.07.052-.043 5.04-5.037a1 1 0 1 0-1.415-1.414L9 11.583V3a2 2 0 0 0-2-2H2a1 1 0 0 0-1 1Z"/></symbol><symbol id="icon-eds-i-arrow-bend-up-medium" viewBox="0 0 24 24"><path d="m11.852 3.011.058-.007L12 3l.075.003.126.017.111.03.111.044.098.052.104.074.082.073 6 6a1 1 0 1 1-1.414 1.414L13 6.415v12.2C13 19.925 11.964 21 10.667 21H4a1 1 0 0 1 0-2h6.667c.175 0 .333-.164.333-.385v-12.2l-4.293 4.292a1 1 0 0 1-1.32.083l-.094-.083a1 1 0 0 1 0-1.414l6-6c.035-.036.073-.068.112-.097l.11-.071.114-.054.105-.035.118-.025Z"/></symbol><symbol id="icon-eds-i-arrow-bend-up-small" viewBox="0 0 16 16"><path d="M1 13.998a1 1 0 0 1 1-1h5V4.413L3.707 7.705a1 1 0 0 1-1.32.084l-.094-.084a1 1 0 0 1 0-1.414l5-5 .063-.059.093-.068.081-.05.105-.047.104-.035.105-.022L7.94 1l.136.001.122.017.113.03.103.04.1.053.102.07.052.043 5.04 5.037a1 1 0 1 1-1.415 1.414L9 4.415v8.583a2 2 0 0 1-2 2H2a1 1 0 0 1-1-1Z"/></symbol><symbol id="icon-eds-i-arrow-diagonal-medium" viewBox="0 0 24 24"><path d="M14 3h6l.075.003.126.017.111.03.111.044.098.052.096.067.09.08c.036.035.068.073.097.112l.071.11.054.114.035.105.03.148L21 4v6a1 1 0 0 1-2 0V6.414l-4.293 4.293a1 1 0 0 1-1.414-1.414L17.584 5H14a1 1 0 0 1-.993-.883L13 4a1 1 0 0 1 1-1ZM4 13a1 1 0 0 1 1 1v3.584l4.293-4.291a1 1 0 1 1 1.414 1.414L6.414 19H10a1 1 0 0 1 .993.883L11 20a1 1 0 0 1-1 1l-6.075-.003-.126-.017-.111-.03-.111-.044-.098-.052-.096-.067-.09-.08a1.01 1.01 0 0 1-.097-.112l-.071-.11-.054-.114-.035-.105-.025-.118-.007-.058L3 20v-6a1 1 0 0 1 1-1Z"/></symbol><symbol id="icon-eds-i-arrow-diagonal-small" viewBox="0 0 16 16"><path d="m2 15-.082-.004-.119-.016-.111-.03-.111-.044-.098-.052-.096-.067-.09-.08a1.008 1.008 0 0 1-.097-.112l-.071-.11-.031-.062-.034-.081-.024-.076-.025-.118-.007-.058L1 14.02V9a1 1 0 1 1 2 0v2.584l2.793-2.791a1 1 0 1 1 1.414 1.414L4.414 13H7a1 1 0 0 1 .993.883L8 14a1 1 0 0 1-1 1H2ZM14 1l.081.003.12.017.111.03.111.044.098.052.096.067.09.08c.036.035.068.073.097.112l.071.11.031.062.034.081.024.076.03.148L15 2v5a1 1 0 0 1-2 0V4.414l-2.96 2.96A1 1 0 1 1 8.626 5.96L11.584 3H9a1 1 0 0 1-.993-.883L8 2a1 1 0 0 1 1-1h5Z"/></symbol><symbol id="icon-eds-i-arrow-down-medium" viewBox="0 0 24 24"><path d="m20.707 12.728-7.99 7.98a.996.996 0 0 1-.561.281l-.157.011a.998.998 0 0 1-.788-.384l-7.918-7.908a1 1 0 0 1 1.414-1.416L11 17.576V4a1 1 0 0 1 2 0v13.598l6.293-6.285a1 1 0 0 1 1.32-.082l.095.083a1 1 0 0 1-.001 1.414Z"/></symbol><symbol id="icon-eds-i-arrow-down-small" viewBox="0 0 16 16"><path d="m1.293 8.707 6 6 .063.059.093.069.081.048.105.049.104.034.056.013.118.017L8 15l.076-.003.122-.017.113-.03.085-.032.063-.03.098-.058.06-.043.05-.043 6.04-6.037a1 1 0 0 0-1.414-1.414L9 11.583V2a1 1 0 1 0-2 0v9.585L2.707 7.293a1 1 0 0 0-1.32-.083l-.094.083a1 1 0 0 0 0 1.414Z"/></symbol><symbol id="icon-eds-i-arrow-left-medium" viewBox="0 0 24 24"><path d="m11.272 3.293-7.98 7.99a.996.996 0 0 0-.281.561L3 12.001c0 .32.15.605.384.788l7.908 7.918a1 1 0 0 0 1.416-1.414L6.424 13H20a1 1 0 0 0 0-2H6.402l6.285-6.293a1 1 0 0 0 .082-1.32l-.083-.095a1 1 0 0 0-1.414.001Z"/></symbol><symbol id="icon-eds-i-arrow-left-small" viewBox="0 0 16 16"><path d="m7.293 1.293-6 6-.059.063-.069.093-.048.081-.049.105-.034.104-.013.056-.017.118L1 8l.003.076.017.122.03.113.032.085.03.063.058.098.043.06.043.05 6.037 6.04a1 1 0 0 0 1.414-1.414L4.417 9H14a1 1 0 0 0 0-2H4.415l4.292-4.293a1 1 0 0 0 .083-1.32l-.083-.094a1 1 0 0 0-1.414 0Z"/></symbol><symbol id="icon-eds-i-arrow-right-medium" viewBox="0 0 24 24"><path d="m12.728 3.293 7.98 7.99a.996.996 0 0 1 .281.561l.011.157c0 .32-.15.605-.384.788l-7.908 7.918a1 1 0 0 1-1.416-1.414L17.576 13H4a1 1 0 0 1 0-2h13.598l-6.285-6.293a1 1 0 0 1-.082-1.32l.083-.095a1 1 0 0 1 1.414.001Z"/></symbol><symbol id="icon-eds-i-arrow-right-small" viewBox="0 0 16 16"><path d="m8.707 1.293 6 6 .059.063.069.093.048.081.049.105.034.104.013.056.017.118L15 8l-.003.076-.017.122-.03.113-.032.085-.03.063-.058.098-.043.06-.043.05-6.037 6.04a1 1 0 0 1-1.414-1.414L11.583 9H2a1 1 0 1 1 0-2h9.585L7.293 2.707a1 1 0 0 1-.083-1.32l.083-.094a1 1 0 0 1 1.414 0Z"/></symbol><symbol id="icon-eds-i-arrow-up-medium" viewBox="0 0 24 24"><path d="m3.293 11.272 7.99-7.98a.996.996 0 0 1 .561-.281L12.001 3c.32 0 .605.15.788.384l7.918 7.908a1 1 0 0 1-1.414 1.416L13 6.424V20a1 1 0 0 1-2 0V6.402l-6.293 6.285a1 1 0 0 1-1.32.082l-.095-.083a1 1 0 0 1 .001-1.414Z"/></symbol><symbol id="icon-eds-i-arrow-up-small" viewBox="0 0 16 16"><path d="m1.293 7.293 6-6 .063-.059.093-.069.081-.048.105-.049.104-.034.056-.013.118-.017L8 1l.076.003.122.017.113.03.085.032.063.03.098.058.06.043.05.043 6.04 6.037a1 1 0 0 1-1.414 1.414L9 4.417V14a1 1 0 0 1-2 0V4.415L2.707 8.707a1 1 0 0 1-1.32.083l-.094-.083a1 1 0 0 1 0-1.414Z"/></symbol><symbol id="icon-eds-i-article-medium" viewBox="0 0 24 24"><path d="M8 7a1 1 0 0 0 0 2h4a1 1 0 1 0 0-2H8ZM8 11a1 1 0 1 0 0 2h8a1 1 0 1 0 0-2H8ZM7 16a1 1 0 0 1 1-1h8a1 1 0 1 1 0 2H8a1 1 0 0 1-1-1Z"/><path d="M5.545 1A2.542 2.542 0 0 0 3 3.538v16.924A2.542 2.542 0 0 0 5.545 23h12.91A2.542 2.542 0 0 0 21 20.462V3.5A2.5 2.5 0 0 0 18.5 1H5.545ZM5 3.538C5 3.245 5.24 3 5.545 3H18.5a.5.5 0 0 1 .5.5v16.962c0 .293-.24.538-.546.538H5.545A.542.542 0 0 1 5 20.462V3.538Z" clip-rule="evenodd"/></symbol><symbol id="icon-eds-i-book-medium" viewBox="0 0 24 24"><path d="M18.5 1A2.5 2.5 0 0 1 21 3.5v12c0 1.16-.79 2.135-1.86 2.418l-.14.031V21h1a1 1 0 0 1 .993.883L21 22a1 1 0 0 1-1 1H6.5A3.5 3.5 0 0 1 3 19.5v-15A3.5 3.5 0 0 1 6.5 1h12ZM17 18H6.5a1.5 1.5 0 0 0-1.493 1.356L5 19.5A1.5 1.5 0 0 0 6.5 21H17v-3Zm1.5-15h-12A1.5 1.5 0 0 0 5 4.5v11.837l.054-.025a3.481 3.481 0 0 1 1.254-.307L6.5 16h12a.5.5 0 0 0 .492-.41L19 15.5v-12a.5.5 0 0 0-.5-.5ZM15 6a1 1 0 0 1 0 2H9a1 1 0 1 1 0-2h6Z"/></symbol><symbol id="icon-eds-i-book-series-medium" viewBox="0 0 24 24"><path fill-rule="evenodd" d="M1 3.786C1 2.759 1.857 2 2.82 2H6.18c.964 0 1.82.759 1.82 1.786V4h3.168c.668 0 1.298.364 1.616.938.158-.109.333-.195.523-.252l3.216-.965c.923-.277 1.962.204 2.257 1.187l4.146 13.82c.296.984-.307 1.957-1.23 2.234l-3.217.965c-.923.277-1.962-.203-2.257-1.187L13 10.005v10.21c0 1.04-.878 1.785-1.834 1.785H7.833c-.291 0-.575-.07-.83-.195A1.849 1.849 0 0 1 6.18 22H2.821C1.857 22 1 21.241 1 20.214V3.786ZM3 4v11h3V4H3Zm0 16v-3h3v3H3Zm15.075-.04-.814-2.712 2.874-.862.813 2.712-2.873.862Zm1.485-5.49-2.874.862-2.634-8.782 2.873-.862 2.635 8.782ZM8 20V6h3v14H8Z" clip-rule="evenodd"/></symbol><symbol id="icon-eds-i-calendar-acceptance-medium" viewBox="0 0 24 24"><path d="M17 2a1 1 0 0 1 1 1v1h1.5C20.817 4 22 5.183 22 6.5v13c0 1.317-1.183 2.5-2.5 2.5h-15C3.183 22 2 20.817 2 19.5v-13C2 5.183 3.183 4 4.5 4a1 1 0 1 1 0 2c-.212 0-.5.288-.5.5v13c0 .212.288.5.5.5h15c.212 0 .5-.288.5-.5v-13c0-.212-.288-.5-.5-.5H18v1a1 1 0 0 1-2 0V3a1 1 0 0 1 1-1Zm-.534 7.747a1 1 0 0 1 .094 1.412l-4.846 5.538a1 1 0 0 1-1.352.141l-2.77-2.076a1 1 0 0 1 1.2-1.6l2.027 1.519 4.236-4.84a1 1 0 0 1 1.411-.094ZM7.5 2a1 1 0 0 1 1 1v1H14a1 1 0 0 1 0 2H8.5v1a1 1 0 1 1-2 0V3a1 1 0 0 1 1-1Z"/></symbol><symbol id="icon-eds-i-calendar-date-medium" viewBox="0 0 24 24"><path d="M17 2a1 1 0 0 1 1 1v1h1.5C20.817 4 22 5.183 22 6.5v13c0 1.317-1.183 2.5-2.5 2.5h-15C3.183 22 2 20.817 2 19.5v-13C2 5.183 3.183 4 4.5 4a1 1 0 1 1 0 2c-.212 0-.5.288-.5.5v13c0 .212.288.5.5.5h15c.212 0 .5-.288.5-.5v-13c0-.212-.288-.5-.5-.5H18v1a1 1 0 0 1-2 0V3a1 1 0 0 1 1-1ZM8 15a1 1 0 1 1 0 2 1 1 0 0 1 0-2Zm4 0a1 1 0 1 1 0 2 1 1 0 0 1 0-2Zm-4-4a1 1 0 1 1 0 2 1 1 0 0 1 0-2Zm4 0a1 1 0 1 1 0 2 1 1 0 0 1 0-2Zm4 0a1 1 0 1 1 0 2 1 1 0 0 1 0-2ZM7.5 2a1 1 0 0 1 1 1v1H14a1 1 0 0 1 0 2H8.5v1a1 1 0 1 1-2 0V3a1 1 0 0 1 1-1Z"/></symbol><symbol id="icon-eds-i-calendar-decision-medium" viewBox="0 0 24 24"><path d="M17 2a1 1 0 0 1 1 1v1h1.5C20.817 4 22 5.183 22 6.5v13c0 1.317-1.183 2.5-2.5 2.5h-15C3.183 22 2 20.817 2 19.5v-13C2 5.183 3.183 4 4.5 4a1 1 0 1 1 0 2c-.212 0-.5.288-.5.5v13c0 .212.288.5.5.5h15c.212 0 .5-.288.5-.5v-13c0-.212-.288-.5-.5-.5H18v1a1 1 0 0 1-2 0V3a1 1 0 0 1 1-1Zm-2.935 8.246 2.686 2.645c.34.335.34.883 0 1.218l-2.686 2.645a.858.858 0 0 1-1.213-.009.854.854 0 0 1 .009-1.21l1.05-1.035H7.984a.992.992 0 0 1-.984-1c0-.552.44-1 .984-1h5.928l-1.051-1.036a.854.854 0 0 1-.085-1.121l.076-.088a.858.858 0 0 1 1.213-.009ZM7.5 2a1 1 0 0 1 1 1v1H14a1 1 0 0 1 0 2H8.5v1a1 1 0 1 1-2 0V3a1 1 0 0 1 1-1Z"/></symbol><symbol id="icon-eds-i-calendar-impact-factor-medium" viewBox="0 0 24 24"><path d="M17 2a1 1 0 0 1 1 1v1h1.5C20.817 4 22 5.183 22 6.5v13c0 1.317-1.183 2.5-2.5 2.5h-15C3.183 22 2 20.817 2 19.5v-13C2 5.183 3.183 4 4.5 4a1 1 0 1 1 0 2c-.212 0-.5.288-.5.5v13c0 .212.288.5.5.5h15c.212 0 .5-.288.5-.5v-13c0-.212-.288-.5-.5-.5H18v1a1 1 0 0 1-2 0V3a1 1 0 0 1 1-1Zm-3.2 6.924a.48.48 0 0 1 .125.544l-1.52 3.283h2.304c.27 0 .491.215.491.483a.477.477 0 0 1-.13.327l-4.18 4.484a.498.498 0 0 1-.69.031.48.48 0 0 1-.125-.544l1.52-3.284H9.291a.487.487 0 0 1-.491-.482c0-.121.047-.238.13-.327l4.18-4.484a.498.498 0 0 1 .69-.031ZM7.5 2a1 1 0 0 1 1 1v1H14a1 1 0 0 1 0 2H8.5v1a1 1 0 1 1-2 0V3a1 1 0 0 1 1-1Z"/></symbol><symbol id="icon-eds-i-call-papers-medium" viewBox="0 0 24 24"><g><path d="m20.707 2.883-1.414 1.414a1 1 0 0 0 1.414 1.414l1.414-1.414a1 1 0 0 0-1.414-1.414Z"/><path d="M6 16.054c0 2.026 1.052 2.943 3 2.943a1 1 0 1 1 0 2c-2.996 0-5-1.746-5-4.943v-1.227a4.068 4.068 0 0 1-1.83-1.189 4.553 4.553 0 0 1-.87-1.455 4.868 4.868 0 0 1-.3-1.686c0-1.17.417-2.298 1.17-3.14.38-.426.834-.767 1.338-1 .51-.237 1.06-.36 1.617-.36L6.632 6H7l7.932-2.895A2.363 2.363 0 0 1 18 5.36v9.28a2.36 2.36 0 0 1-3.069 2.25l.084.03L7 14.997H6v1.057Zm9.637-11.057a.415.415 0 0 0-.083.008L8 7.638v5.536l7.424 1.786.104.02c.035.01.072.02.109.02.2 0 .363-.16.363-.36V5.36c0-.2-.163-.363-.363-.363Zm-9.638 3h-.874a1.82 1.82 0 0 0-.625.111l-.15.063a2.128 2.128 0 0 0-.689.517c-.42.47-.661 1.123-.661 1.81 0 .34.06.678.176.992.114.308.28.585.485.816.4.447.925.691 1.464.691h.874v-5Z" clip-rule="evenodd"/><path d="M20 8.997h2a1 1 0 1 1 0 2h-2a1 1 0 1 1 0-2ZM20.707 14.293l1.414 1.414a1 1 0 0 1-1.414 1.414l-1.414-1.414a1 1 0 0 1 1.414-1.414Z"/></g></symbol><symbol id="icon-eds-i-card-medium" viewBox="0 0 24 24"><path d="M19.615 2c.315 0 .716.067 1.14.279.76.38 1.245 1.107 1.245 2.106v15.23c0 .315-.067.716-.279 1.14-.38.76-1.107 1.245-2.106 1.245H4.385a2.56 2.56 0 0 1-1.14-.279C2.485 21.341 2 20.614 2 19.615V4.385c0-.315.067-.716.279-1.14C2.659 2.485 3.386 2 4.385 2h15.23Zm0 2H4.385c-.213 0-.265.034-.317.14A.71.71 0 0 0 4 4.385v15.23c0 .213.034.265.14.317a.71.71 0 0 0 .245.068h15.23c.213 0 .265-.034.317-.14a.71.71 0 0 0 .068-.245V4.385c0-.213-.034-.265-.14-.317A.71.71 0 0 0 19.615 4ZM17 16a1 1 0 0 1 0 2H7a1 1 0 0 1 0-2h10Zm0-3a1 1 0 0 1 0 2H7a1 1 0 0 1 0-2h10Zm-.5-7A1.5 1.5 0 0 1 18 7.5v3a1.5 1.5 0 0 1-1.5 1.5h-9A1.5 1.5 0 0 1 6 10.5v-3A1.5 1.5 0 0 1 7.5 6h9ZM16 8H8v2h8V8Z"/></symbol><symbol id="icon-eds-i-cart-medium" viewBox="0 0 24 24"><path d="M5.76 1a1 1 0 0 1 .994.902L7.155 6h13.34c.18 0 .358.02.532.057l.174.045a2.5 2.5 0 0 1 1.693 3.103l-2.069 7.03c-.36 1.099-1.398 1.823-2.49 1.763H8.65c-1.272.015-2.352-.927-2.546-2.244L4.852 3H2a1 1 0 0 1-.993-.883L1 2a1 1 0 0 1 1-1h3.76Zm2.328 14.51a.555.555 0 0 0 .55.488l9.751.001a.533.533 0 0 0 .527-.357l2.059-7a.5.5 0 0 0-.48-.642H7.351l.737 7.51ZM18 19a2 2 0 1 1 0 4 2 2 0 0 1 0-4ZM8 19a2 2 0 1 1 0 4 2 2 0 0 1 0-4Z"/></symbol><symbol id="icon-eds-i-check-circle-medium" viewBox="0 0 24 24"><path d="M12 1c6.075 0 11 4.925 11 11s-4.925 11-11 11S1 18.075 1 12 5.925 1 12 1Zm0 2a9 9 0 1 0 0 18 9 9 0 0 0 0-18Zm5.125 4.72a1 1 0 0 1 .156 1.405l-6 7.5a1 1 0 0 1-1.421.143l-3-2.5a1 1 0 0 1 1.28-1.536l2.217 1.846 5.362-6.703a1 1 0 0 1 1.406-.156Z"/></symbol><symbol id="icon-eds-i-check-filled-medium" viewBox="0 0 24 24"><path d="M12 1c6.075 0 11 4.925 11 11s-4.925 11-11 11S1 18.075 1 12 5.925 1 12 1Zm5.125 6.72a1 1 0 0 0-1.406.155l-5.362 6.703-2.217-1.846a1 1 0 1 0-1.28 1.536l3 2.5a1 1 0 0 0 1.42-.143l6-7.5a1 1 0 0 0-.155-1.406Z"/></symbol><symbol id="icon-eds-i-chevron-down-medium" viewBox="0 0 24 24"><path d="M3.305 8.28a1 1 0 0 0-.024 1.415l7.495 7.762c.314.345.757.543 1.224.543.467 0 .91-.198 1.204-.522l7.515-7.783a1 1 0 1 0-1.438-1.39L12 15.845l-7.28-7.54A1 1 0 0 0 3.4 8.2l-.096.082Z"/></symbol><symbol id="icon-eds-i-chevron-down-small" viewBox="0 0 16 16"><path d="M13.692 5.278a1 1 0 0 1 .03 1.414L9.103 11.51a1.491 1.491 0 0 1-2.188.019L2.278 6.692a1 1 0 0 1 1.444-1.384L8 9.771l4.278-4.463a1 1 0 0 1 1.318-.111l.096.081Z"/></symbol><symbol id="icon-eds-i-chevron-left-medium" viewBox="0 0 24 24"><path d="M15.72 3.305a1 1 0 0 0-1.415-.024l-7.762 7.495A1.655 1.655 0 0 0 6 12c0 .467.198.91.522 1.204l7.783 7.515a1 1 0 1 0 1.39-1.438L8.155 12l7.54-7.28A1 1 0 0 0 15.8 3.4l-.082-.096Z"/></symbol><symbol id="icon-eds-i-chevron-left-small" viewBox="0 0 16 16"><path d="M10.722 2.308a1 1 0 0 0-1.414-.03L4.49 6.897a1.491 1.491 0 0 0-.019 2.188l4.838 4.637a1 1 0 1 0 1.384-1.444L6.229 8l4.463-4.278a1 1 0 0 0 .111-1.318l-.081-.096Z"/></symbol><symbol id="icon-eds-i-chevron-right-medium" viewBox="0 0 24 24"><path d="M8.28 3.305a1 1 0 0 1 1.415-.024l7.762 7.495c.345.314.543.757.543 1.224 0 .467-.198.91-.522 1.204l-7.783 7.515a1 1 0 1 1-1.39-1.438L15.845 12l-7.54-7.28A1 1 0 0 1 8.2 3.4l.082-.096Z"/></symbol><symbol id="icon-eds-i-chevron-right-small" viewBox="0 0 16 16"><path d="M5.278 2.308a1 1 0 0 1 1.414-.03l4.819 4.619a1.491 1.491 0 0 1 .019 2.188l-4.838 4.637a1 1 0 1 1-1.384-1.444L9.771 8 5.308 3.722a1 1 0 0 1-.111-1.318l.081-.096Z"/></symbol><symbol id="icon-eds-i-chevron-up-medium" viewBox="0 0 24 24"><path d="M20.695 15.72a1 1 0 0 0 .024-1.415l-7.495-7.762A1.655 1.655 0 0 0 12 6c-.467 0-.91.198-1.204.522l-7.515 7.783a1 1 0 1 0 1.438 1.39L12 8.155l7.28 7.54a1 1 0 0 0 1.319.106l.096-.082Z"/></symbol><symbol id="icon-eds-i-chevron-up-small" viewBox="0 0 16 16"><path d="M13.692 10.722a1 1 0 0 0 .03-1.414L9.103 4.49a1.491 1.491 0 0 0-2.188-.019L2.278 9.308a1 1 0 0 0 1.444 1.384L8 6.229l4.278 4.463a1 1 0 0 0 1.318.111l.096-.081Z"/></symbol><symbol id="icon-eds-i-citations-medium" viewBox="0 0 24 24"><path d="M15.59 1a1 1 0 0 1 .706.291l5.41 5.385a1 1 0 0 1 .294.709v13.077c0 .674-.269 1.32-.747 1.796a2.549 2.549 0 0 1-1.798.742h-5.843a1 1 0 1 1 0-2h5.843a.549.549 0 0 0 .387-.16.535.535 0 0 0 .158-.378V7.8L15.178 3H5.545a.543.543 0 0 0-.538.451L5 3.538v8.607a1 1 0 0 1-2 0V3.538A2.542 2.542 0 0 1 5.545 1h10.046ZM5.483 14.35c.197.26.17.62-.049.848l-.095.083-.016.011c-.36.24-.628.45-.804.634-.393.409-.59.93-.59 1.562.077-.019.192-.028.345-.028.442 0 .84.158 1.195.474.355.316.532.716.532 1.2 0 .501-.173.9-.518 1.198-.345.298-.767.446-1.266.446-.672 0-1.209-.195-1.612-.585-.403-.39-.604-.976-.604-1.757 0-.744.11-1.39.33-1.938.222-.549.49-1.009.807-1.38a4.28 4.28 0 0 1 .992-.88c.07-.043.148-.087.232-.133a.881.881 0 0 1 1.121.245Zm5 0c.197.26.17.62-.049.848l-.095.083-.016.011c-.36.24-.628.45-.804.634-.393.409-.59.93-.59 1.562.077-.019.192-.028.345-.028.442 0 .84.158 1.195.474.355.316.532.716.532 1.2 0 .501-.173.9-.518 1.198-.345.298-.767.446-1.266.446-.672 0-1.209-.195-1.612-.585-.403-.39-.604-.976-.604-1.757 0-.744.11-1.39.33-1.938.222-.549.49-1.009.807-1.38a4.28 4.28 0 0 1 .992-.88c.07-.043.148-.087.232-.133a.881.881 0 0 1 1.121.245Z"/></symbol><symbol id="icon-eds-i-clipboard-check-medium" viewBox="0 0 24 24"><path d="M14.4 1c1.238 0 2.274.865 2.536 2.024L18.5 3C19.886 3 21 4.14 21 5.535v14.93C21 21.86 19.886 23 18.5 23h-13C4.114 23 3 21.86 3 20.465V5.535C3 4.14 4.114 3 5.5 3h1.57c.27-1.147 1.3-2 2.53-2h4.8Zm4.115 4-1.59.024A2.601 2.601 0 0 1 14.4 7H9.6c-1.23 0-2.26-.853-2.53-2H5.5c-.27 0-.5.234-.5.535v14.93c0 .3.23.535.5.535h13c.27 0 .5-.234.5-.535V5.535c0-.3-.23-.535-.485-.535Zm-1.909 4.205a1 1 0 0 1 .19 1.401l-5.334 7a1 1 0 0 1-1.344.23l-2.667-1.75a1 1 0 1 1 1.098-1.672l1.887 1.238 4.769-6.258a1 1 0 0 1 1.401-.19ZM14.4 3H9.6a.6.6 0 0 0-.6.6v.8a.6.6 0 0 0 .6.6h4.8a.6.6 0 0 0 .6-.6v-.8a.6.6 0 0 0-.6-.6Z"/></symbol><symbol id="icon-eds-i-clipboard-report-medium" viewBox="0 0 24 24"><path d="M14.4 1c1.238 0 2.274.865 2.536 2.024L18.5 3C19.886 3 21 4.14 21 5.535v14.93C21 21.86 19.886 23 18.5 23h-13C4.114 23 3 21.86 3 20.465V5.535C3 4.14 4.114 3 5.5 3h1.57c.27-1.147 1.3-2 2.53-2h4.8Zm4.115 4-1.59.024A2.601 2.601 0 0 1 14.4 7H9.6c-1.23 0-2.26-.853-2.53-2H5.5c-.27 0-.5.234-.5.535v14.93c0 .3.23.535.5.535h13c.27 0 .5-.234.5-.535V5.535c0-.3-.23-.535-.485-.535Zm-2.658 10.929a1 1 0 0 1 0 2H8a1 1 0 0 1 0-2h7.857Zm0-3.929a1 1 0 0 1 0 2H8a1 1 0 0 1 0-2h7.857ZM14.4 3H9.6a.6.6 0 0 0-.6.6v.8a.6.6 0 0 0 .6.6h4.8a.6.6 0 0 0 .6-.6v-.8a.6.6 0 0 0-.6-.6Z"/></symbol><symbol id="icon-eds-i-close-medium" viewBox="0 0 24 24"><path d="M12 1c6.075 0 11 4.925 11 11s-4.925 11-11 11S1 18.075 1 12 5.925 1 12 1Zm0 2a9 9 0 1 0 0 18 9 9 0 0 0 0-18ZM8.707 7.293 12 10.585l3.293-3.292a1 1 0 0 1 1.414 1.414L13.415 12l3.292 3.293a1 1 0 0 1-1.414 1.414L12 13.415l-3.293 3.292a1 1 0 1 1-1.414-1.414L10.585 12 7.293 8.707a1 1 0 0 1 1.414-1.414Z"/></symbol><symbol id="icon-eds-i-cloud-upload-medium" viewBox="0 0 24 24"><path d="m12.852 10.011.028-.004L13 10l.075.003.126.017.086.022.136.052.098.052.104.074.082.073 3 3a1 1 0 0 1 0 1.414l-.094.083a1 1 0 0 1-1.32-.083L14 13.416V20a1 1 0 0 1-2 0v-6.586l-1.293 1.293a1 1 0 0 1-1.32.083l-.094-.083a1 1 0 0 1 0-1.414l3-3 .112-.097.11-.071.114-.054.105-.035.118-.025Zm.587-7.962c3.065.362 5.497 2.662 5.992 5.562l.013.085.207.073c2.117.782 3.496 2.845 3.337 5.097l-.022.226c-.297 2.561-2.503 4.491-5.124 4.502a1 1 0 1 1-.009-2c1.619-.007 2.967-1.186 3.147-2.733.179-1.542-.86-2.979-2.487-3.353-.512-.149-.894-.579-.981-1.165-.21-2.237-2-4.035-4.308-4.308-2.31-.273-4.497 1.06-5.25 3.19l-.049.113c-.234.468-.718.756-1.176.743-1.418.057-2.689.857-3.32 2.084a3.668 3.668 0 0 0 .262 3.798c.796 1.136 2.169 1.764 3.583 1.635a1 1 0 1 1 .182 1.992c-2.125.194-4.193-.753-5.403-2.48a5.668 5.668 0 0 1-.403-5.86c.85-1.652 2.449-2.79 4.323-3.092l.287-.039.013-.028c1.207-2.741 4.125-4.404 7.186-4.042Z"/></symbol><symbol id="icon-eds-i-collection-medium" viewBox="0 0 24 24"><path d="M21 7a1 1 0 0 1 1 1v12.5a2.5 2.5 0 0 1-2.5 2.5H8a1 1 0 0 1 0-2h11.5a.5.5 0 0 0 .5-.5V8a1 1 0 0 1 1-1Zm-5.5-5A2.5 2.5 0 0 1 18 4.5v12a2.5 2.5 0 0 1-2.5 2.5h-11A2.5 2.5 0 0 1 2 16.5v-12A2.5 2.5 0 0 1 4.5 2h11Zm0 2h-11a.5.5 0 0 0-.5.5v12a.5.5 0 0 0 .5.5h11a.5.5 0 0 0 .5-.5v-12a.5.5 0 0 0-.5-.5ZM13 13a1 1 0 0 1 0 2H7a1 1 0 0 1 0-2h6Zm0-3.5a1 1 0 0 1 0 2H7a1 1 0 0 1 0-2h6ZM13 6a1 1 0 0 1 0 2H7a1 1 0 1 1 0-2h6Z"/></symbol><symbol id="icon-eds-i-conference-series-medium" viewBox="0 0 24 24"><path fill-rule="evenodd" d="M4.5 2A2.5 2.5 0 0 0 2 4.5v11A2.5 2.5 0 0 0 4.5 18h2.37l-2.534 2.253a1 1 0 0 0 1.328 1.494L9.88 18H11v3a1 1 0 1 0 2 0v-3h1.12l4.216 3.747a1 1 0 0 0 1.328-1.494L17.13 18h2.37a2.5 2.5 0 0 0 2.5-2.5v-11A2.5 2.5 0 0 0 19.5 2h-15ZM20 6V4.5a.5.5 0 0 0-.5-.5h-15a.5.5 0 0 0-.5.5V6h16ZM4 8v7.5a.5.5 0 0 0 .5.5h15a.5.5 0 0 0 .5-.5V8H4Z" clip-rule="evenodd"/></symbol><symbol id="icon-eds-i-delivery-medium" viewBox="0 0 24 24"><path d="M8.51 20.598a3.037 3.037 0 0 1-3.02 0A2.968 2.968 0 0 1 4.161 19L3.5 19A2.5 2.5 0 0 1 1 16.5v-11A2.5 2.5 0 0 1 3.5 3h10a2.5 2.5 0 0 1 2.45 2.004L16 5h2.527c.976 0 1.855.585 2.27 1.49l2.112 4.62a1 1 0 0 1 .091.416v4.856C23 17.814 21.889 19 20.484 19h-.523a1.01 1.01 0 0 1-.121-.007 2.96 2.96 0 0 1-1.33 1.605 3.037 3.037 0 0 1-3.02 0A2.968 2.968 0 0 1 14.161 19H9.838a2.968 2.968 0 0 1-1.327 1.597Zm-2.024-3.462a.955.955 0 0 0-.481.73L5.999 18l.001.022a.944.944 0 0 0 .388.777l.098.065c.316.181.712.181 1.028 0A.97.97 0 0 0 8 17.978a.95.95 0 0 0-.486-.842 1.037 1.037 0 0 0-1.028 0Zm10 0a.955.955 0 0 0-.481.73l-.005.156a.944.944 0 0 0 .388.777l.098.065c.316.181.712.181 1.028 0a.97.97 0 0 0 .486-.886.95.95 0 0 0-.486-.842 1.037 1.037 0 0 0-1.028 0ZM21 12h-5v3.17a3.038 3.038 0 0 1 2.51.232 2.993 2.993 0 0 1 1.277 1.45l.058.155.058-.005.581-.002c.27 0 .516-.263.516-.618V12Zm-7.5-7h-10a.5.5 0 0 0-.5.5v11a.5.5 0 0 0 .5.5h.662a2.964 2.964 0 0 1 1.155-1.491l.172-.107a3.037 3.037 0 0 1 3.022 0A2.987 2.987 0 0 1 9.843 17H13.5a.5.5 0 0 0 .5-.5v-11a.5.5 0 0 0-.5-.5Zm5.027 2H16v3h4.203l-1.224-2.677a.532.532 0 0 0-.375-.316L18.527 7Z"/></symbol><symbol id="icon-eds-i-download-medium" viewBox="0 0 24 24"><path d="M22 18.5a3.5 3.5 0 0 1-3.5 3.5h-13A3.5 3.5 0 0 1 2 18.5V18a1 1 0 0 1 2 0v.5A1.5 1.5 0 0 0 5.5 20h13a1.5 1.5 0 0 0 1.5-1.5V18a1 1 0 0 1 2 0v.5Zm-3.293-7.793-6 6-.063.059-.093.069-.081.048-.105.049-.104.034-.056.013-.118.017L12 17l-.076-.003-.122-.017-.113-.03-.085-.032-.063-.03-.098-.058-.06-.043-.05-.043-6.04-6.037a1 1 0 0 1 1.414-1.414l4.294 4.29L11 3a1 1 0 0 1 2 0l.001 10.585 4.292-4.292a1 1 0 0 1 1.32-.083l.094.083a1 1 0 0 1 0 1.414Z"/></symbol><symbol id="icon-eds-i-edit-medium" viewBox="0 0 24 24"><path d="M17.149 2a2.38 2.38 0 0 1 1.699.711l2.446 2.46a2.384 2.384 0 0 1 .005 3.38L10.01 19.906a1 1 0 0 1-.434.257l-6.3 1.8a1 1 0 0 1-1.237-1.237l1.8-6.3a1 1 0 0 1 .257-.434L15.443 2.718A2.385 2.385 0 0 1 17.15 2Zm-3.874 5.689-7.586 7.536-1.234 4.319 4.318-1.234 7.54-7.582-3.038-3.039ZM17.149 4a.395.395 0 0 0-.286.126L14.695 6.28l3.029 3.029 2.162-2.173a.384.384 0 0 0 .106-.197L20 6.864c0-.103-.04-.2-.119-.278l-2.457-2.47A.385.385 0 0 0 17.149 4Z"/></symbol><symbol id="icon-eds-i-education-medium" viewBox="0 0 24 24"><path fill-rule="evenodd" d="M12.41 2.088a1 1 0 0 0-.82 0l-10 4.5a1 1 0 0 0 0 1.824L3 9.047v7.124A3.001 3.001 0 0 0 4 22a3 3 0 0 0 1-5.83V9.948l1 .45V14.5a1 1 0 0 0 .087.408L7 14.5c-.913.408-.912.41-.912.41l.001.003.003.006.007.015a1.988 1.988 0 0 0 .083.16c.054.097.131.225.236.373.21.297.53.68.993 1.057C8.351 17.292 9.824 18 12 18c2.176 0 3.65-.707 4.589-1.476.463-.378.783-.76.993-1.057a4.162 4.162 0 0 0 .319-.533l.007-.015.003-.006v-.003h.002s0-.002-.913-.41l.913.408A1 1 0 0 0 18 14.5v-4.103l4.41-1.985a1 1 0 0 0 0-1.824l-10-4.5ZM16 11.297l-3.59 1.615a1 1 0 0 1-.82 0L8 11.297v2.94a3.388 3.388 0 0 0 .677.739C9.267 15.457 10.294 16 12 16s2.734-.543 3.323-1.024a3.388 3.388 0 0 0 .677-.739v-2.94ZM4.437 7.5 12 4.097 19.563 7.5 12 10.903 4.437 7.5ZM3 19a1 1 0 1 1 2 0 1 1 0 0 1-2 0Z" clip-rule="evenodd"/></symbol><symbol id="icon-eds-i-error-diamond-medium" viewBox="0 0 24 24"><path d="M12.002 1c.702 0 1.375.279 1.871.775l8.35 8.353a2.646 2.646 0 0 1 .001 3.744l-8.353 8.353a2.646 2.646 0 0 1-3.742 0l-8.353-8.353a2.646 2.646 0 0 1 0-3.744l8.353-8.353.156-.142c.424-.362.952-.58 1.507-.625l.21-.008Zm0 2a.646.646 0 0 0-.38.123l-.093.08-8.34 8.34a.646.646 0 0 0-.18.355L3 12c0 .171.068.336.19.457l8.353 8.354a.646.646 0 0 0 .914 0l8.354-8.354a.646.646 0 0 0-.001-.914l-8.351-8.354A.646.646 0 0 0 12.002 3ZM12 14.5a1.5 1.5 0 0 1 .144 2.993L12 17.5a1.5 1.5 0 0 1 0-3ZM12 6a1 1 0 0 1 1 1v5a1 1 0 0 1-2 0V7a1 1 0 0 1 1-1Z"/></symbol><symbol id="icon-eds-i-error-filled-medium" viewBox="0 0 24 24"><path d="M12.002 1c.702 0 1.375.279 1.871.775l8.35 8.353a2.646 2.646 0 0 1 .001 3.744l-8.353 8.353a2.646 2.646 0 0 1-3.742 0l-8.353-8.353a2.646 2.646 0 0 1 0-3.744l8.353-8.353.156-.142c.424-.362.952-.58 1.507-.625l.21-.008ZM12 14.5a1.5 1.5 0 0 0 0 3l.144-.007A1.5 1.5 0 0 0 12 14.5ZM12 6a1 1 0 0 0-1 1v5a1 1 0 0 0 2 0V7a1 1 0 0 0-1-1Z"/></symbol><symbol id="icon-eds-i-external-link-medium" viewBox="0 0 24 24"><path d="M9 2a1 1 0 1 1 0 2H4.6c-.371 0-.6.209-.6.5v15c0 .291.229.5.6.5h14.8c.371 0 .6-.209.6-.5V15a1 1 0 0 1 2 0v4.5c0 1.438-1.162 2.5-2.6 2.5H4.6C3.162 22 2 20.938 2 19.5v-15C2 3.062 3.162 2 4.6 2H9Zm6 0h6l.075.003.126.017.111.03.111.044.098.052.096.067.09.08c.036.035.068.073.097.112l.071.11.054.114.035.105.03.148L22 3v6a1 1 0 0 1-2 0V5.414l-6.693 6.693a1 1 0 0 1-1.414-1.414L18.584 4H15a1 1 0 0 1-.993-.883L14 3a1 1 0 0 1 1-1Z"/></symbol><symbol id="icon-eds-i-external-link-small" viewBox="0 0 16 16"><path d="M5 1a1 1 0 1 1 0 2l-2-.001V13L13 13v-2a1 1 0 0 1 2 0v2c0 1.15-.93 2-2.067 2H3.067C1.93 15 1 14.15 1 13V3c0-1.15.93-2 2.067-2H5Zm4 0h5l.075.003.126.017.111.03.111.044.098.052.096.067.09.08.044.047.073.093.051.083.054.113.035.105.03.148L15 2v5a1 1 0 0 1-2 0V4.414L9.107 8.307a1 1 0 0 1-1.414-1.414L11.584 3H9a1 1 0 0 1-.993-.883L8 2a1 1 0 0 1 1-1Z"/></symbol><symbol id="icon-eds-i-file-download-medium" viewBox="0 0 24 24"><path d="M14.5 1a1 1 0 0 1 .707.293l5.5 5.5A1 1 0 0 1 21 7.5v12.962A2.542 2.542 0 0 1 18.455 23H5.545A2.542 2.542 0 0 1 3 20.462V3.538A2.542 2.542 0 0 1 5.545 1H14.5Zm-.415 2h-8.54A.542.542 0 0 0 5 3.538v16.924c0 .296.243.538.545.538h12.91a.542.542 0 0 0 .545-.538V7.915L14.085 3ZM12 7a1 1 0 0 1 1 1v6.585l2.293-2.292a1 1 0 0 1 1.32-.083l.094.083a1 1 0 0 1 0 1.414l-4 4a1.008 1.008 0 0 1-.112.097l-.11.071-.114.054-.105.035-.149.03L12 18l-.075-.003-.126-.017-.111-.03-.111-.044-.098-.052-.096-.067-.09-.08-4-4a1 1 0 0 1 1.414-1.414L11 14.585V8a1 1 0 0 1 1-1Z"/></symbol><symbol id="icon-eds-i-file-report-medium" viewBox="0 0 24 24"><path d="M14.5 1a1 1 0 0 1 .707.293l5.5 5.5A1 1 0 0 1 21 7.5v12.962c0 .674-.269 1.32-.747 1.796a2.549 2.549 0 0 1-1.798.742H5.545c-.674 0-1.32-.267-1.798-.742A2.535 2.535 0 0 1 3 20.462V3.538A2.542 2.542 0 0 1 5.545 1H14.5Zm-.415 2h-8.54A.542.542 0 0 0 5 3.538v16.924c0 .142.057.278.158.379.102.102.242.159.387.159h12.91a.549.549 0 0 0 .387-.16.535.535 0 0 0 .158-.378V7.915L14.085 3ZM16 17a1 1 0 0 1 0 2H8a1 1 0 0 1 0-2h8Zm0-3a1 1 0 0 1 0 2H8a1 1 0 0 1 0-2h8Zm-4.793-6.207L13 9.585l1.793-1.792a1 1 0 0 1 1.32-.083l.094.083a1 1 0 0 1 0 1.414l-2.5 2.5a1 1 0 0 1-1.414 0L10.5 9.915l-1.793 1.792a1 1 0 0 1-1.32.083l-.094-.083a1 1 0 0 1 0-1.414l2.5-2.5a1 1 0 0 1 1.414 0Z"/></symbol><symbol id="icon-eds-i-file-text-medium" viewBox="0 0 24 24"><path d="M14.5 1a1 1 0 0 1 .707.293l5.5 5.5A1 1 0 0 1 21 7.5v12.962A2.542 2.542 0 0 1 18.455 23H5.545A2.542 2.542 0 0 1 3 20.462V3.538A2.542 2.542 0 0 1 5.545 1H14.5Zm-.415 2h-8.54A.542.542 0 0 0 5 3.538v16.924c0 .296.243.538.545.538h12.91a.542.542 0 0 0 .545-.538V7.915L14.085 3ZM16 15a1 1 0 0 1 0 2H8a1 1 0 0 1 0-2h8Zm0-4a1 1 0 0 1 0 2H8a1 1 0 0 1 0-2h8Zm-5-4a1 1 0 0 1 0 2H8a1 1 0 1 1 0-2h3Z"/></symbol><symbol id="icon-eds-i-file-upload-medium" viewBox="0 0 24 24"><path d="M14.5 1a1 1 0 0 1 .707.293l5.5 5.5A1 1 0 0 1 21 7.5v12.962A2.542 2.542 0 0 1 18.455 23H5.545A2.542 2.542 0 0 1 3 20.462V3.538A2.542 2.542 0 0 1 5.545 1H14.5Zm-.415 2h-8.54A.542.542 0 0 0 5 3.538v16.924c0 .296.243.538.545.538h12.91a.542.542 0 0 0 .545-.538V7.915L14.085 3Zm-2.233 4.011.058-.007L12 7l.075.003.126.017.111.03.111.044.098.052.104.074.082.073 4 4a1 1 0 0 1 0 1.414l-.094.083a1 1 0 0 1-1.32-.083L13 10.415V17a1 1 0 0 1-2 0v-6.585l-2.293 2.292a1 1 0 0 1-1.32.083l-.094-.083a1 1 0 0 1 0-1.414l4-4 .112-.097.11-.071.114-.054.105-.035.118-.025Z"/></symbol><symbol id="icon-eds-i-filter-medium" viewBox="0 0 24 24"><path d="M21 2a1 1 0 0 1 .82 1.573L15 13.314V18a1 1 0 0 1-.31.724l-.09.076-4 3A1 1 0 0 1 9 21v-7.684L2.18 3.573a1 1 0 0 1 .707-1.567L3 2h18Zm-1.921 2H4.92l5.9 8.427a1 1 0 0 1 .172.45L11 13v6l2-1.5V13a1 1 0 0 1 .117-.469l.064-.104L19.079 4Z"/></symbol><symbol id="icon-eds-i-funding-medium" viewBox="0 0 24 24"><path fill-rule="evenodd" d="M23 8A7 7 0 1 0 9 8a7 7 0 0 0 14 0ZM9.006 12.225A4.07 4.07 0 0 0 6.12 11.02H2a.979.979 0 1 0 0 1.958h4.12c.558 0 1.094.222 1.489.617l2.207 2.288c.27.27.27.687.012.944a.656.656 0 0 1-.928 0L7.744 15.67a.98.98 0 0 0-1.386 1.384l1.157 1.158c.535.536 1.244.791 1.946.765l.041.002h6.922c.874 0 1.597.748 1.597 1.688 0 .203-.146.354-.309.354H7.755c-.487 0-.96-.178-1.339-.504L2.64 17.259a.979.979 0 0 0-1.28 1.482L5.137 22c.733.631 1.66.979 2.618.979h9.957c1.26 0 2.267-1.043 2.267-2.312 0-2.006-1.584-3.646-3.555-3.646h-4.529a2.617 2.617 0 0 0-.681-2.509l-2.208-2.287ZM16 3a5 5 0 1 0 0 10 5 5 0 0 0 0-10Zm.979 3.5a.979.979 0 1 0-1.958 0v3a.979.979 0 1 0 1.958 0v-3Z" clip-rule="evenodd"/></symbol><symbol id="icon-eds-i-hashtag-medium" viewBox="0 0 24 24"><path d="M12 1c6.075 0 11 4.925 11 11s-4.925 11-11 11S1 18.075 1 12 5.925 1 12 1Zm0 2a9 9 0 1 0 0 18 9 9 0 0 0 0-18ZM9.52 18.189a1 1 0 1 1-1.964-.378l.437-2.274H6a1 1 0 1 1 0-2h2.378l.592-3.076H6a1 1 0 0 1 0-2h3.354l.51-2.65a1 1 0 1 1 1.964.378l-.437 2.272h3.04l.51-2.65a1 1 0 1 1 1.964.378l-.438 2.272H18a1 1 0 0 1 0 2h-1.917l-.592 3.076H18a1 1 0 0 1 0 2h-2.893l-.51 2.652a1 1 0 1 1-1.964-.378l.437-2.274h-3.04l-.51 2.652Zm.895-4.652h3.04l.591-3.076h-3.04l-.591 3.076Z"/></symbol><symbol id="icon-eds-i-home-medium" viewBox="0 0 24 24"><path d="M5 22a1 1 0 0 1-1-1v-8.586l-1.293 1.293a1 1 0 0 1-1.32.083l-.094-.083a1 1 0 0 1 0-1.414l10-10a1 1 0 0 1 1.414 0l10 10a1 1 0 0 1-1.414 1.414L20 12.415V21a1 1 0 0 1-1 1H5Zm7-17.585-6 5.999V20h5v-4a1 1 0 0 1 2 0v4h5v-9.585l-6-6Z"/></symbol><symbol id="icon-eds-i-image-medium" viewBox="0 0 24 24"><path d="M19.615 2A2.385 2.385 0 0 1 22 4.385v15.23A2.385 2.385 0 0 1 19.615 22H4.385A2.385 2.385 0 0 1 2 19.615V4.385A2.385 2.385 0 0 1 4.385 2h15.23Zm0 2H4.385A.385.385 0 0 0 4 4.385v15.23c0 .213.172.385.385.385h1.244l10.228-8.76a1 1 0 0 1 1.254-.037L20 13.392V4.385A.385.385 0 0 0 19.615 4Zm-3.07 9.283L8.703 20h10.912a.385.385 0 0 0 .385-.385v-3.713l-3.455-2.619ZM9.5 6a3.5 3.5 0 1 1 0 7 3.5 3.5 0 0 1 0-7Zm0 2a1.5 1.5 0 1 0 0 3 1.5 1.5 0 0 0 0-3Z"/></symbol><symbol id="icon-eds-i-impact-factor-medium" viewBox="0 0 24 24"><path d="M16.49 2.672c.74.694.986 1.765.632 2.712l-.04.1-1.549 3.54h1.477a2.496 2.496 0 0 1 2.485 2.34l.005.163c0 .618-.23 1.21-.642 1.675l-7.147 7.961a2.48 2.48 0 0 1-3.554.165 2.512 2.512 0 0 1-.633-2.712l.042-.103L9.108 15H7.46c-1.393 0-2.379-1.11-2.455-2.369L5 12.473c0-.593.142-1.145.628-1.692l7.307-7.944a2.48 2.48 0 0 1 3.555-.165ZM14.43 4.164l-7.33 7.97c-.083.093-.101.214-.101.34 0 .277.19.526.46.526h4.163l.097-.009c.015 0 .03.003.046.009.181.078.264.32.186.5l-2.554 5.817a.512.512 0 0 0 .127.552.48.48 0 0 0 .69-.033l7.155-7.97a.513.513 0 0 0 .13-.34.497.497 0 0 0-.49-.502h-3.988a.355.355 0 0 1-.328-.497l2.555-5.844a.512.512 0 0 0-.127-.552.48.48 0 0 0-.69.033Z"/></symbol><symbol id="icon-eds-i-info-circle-medium" viewBox="0 0 24 24"><path d="M12 1c6.075 0 11 4.925 11 11s-4.925 11-11 11S1 18.075 1 12 5.925 1 12 1Zm0 2a9 9 0 1 0 0 18 9 9 0 0 0 0-18Zm0 7a1 1 0 0 1 1 1v5h1.5a1 1 0 0 1 0 2h-5a1 1 0 0 1 0-2H11v-4h-.5a1 1 0 0 1-.993-.883L9.5 11a1 1 0 0 1 1-1H12Zm0-4.5a1.5 1.5 0 0 1 .144 2.993L12 8.5a1.5 1.5 0 0 1 0-3Z"/></symbol><symbol id="icon-eds-i-info-filled-medium" viewBox="0 0 24 24"><path d="M12 1c6.075 0 11 4.925 11 11s-4.925 11-11 11S1 18.075 1 12 5.925 1 12 1Zm0 9h-1.5a1 1 0 0 0-1 1l.007.117A1 1 0 0 0 10.5 12h.5v4H9.5a1 1 0 0 0 0 2h5a1 1 0 0 0 0-2H13v-5a1 1 0 0 0-1-1Zm0-4.5a1.5 1.5 0 0 0 0 3l.144-.007A1.5 1.5 0 0 0 12 5.5Z"/></symbol><symbol id="icon-eds-i-journal-medium" viewBox="0 0 24 24"><path d="M18.5 1A2.5 2.5 0 0 1 21 3.5v14a2.5 2.5 0 0 1-2.5 2.5h-13a.5.5 0 1 0 0 1H20a1 1 0 0 1 0 2H5.5A2.5 2.5 0 0 1 3 20.5v-17A2.5 2.5 0 0 1 5.5 1h13ZM7 3H5.5a.5.5 0 0 0-.5.5v14.549l.016-.002c.104-.02.211-.035.32-.042L5.5 18H7V3Zm11.5 0H9v15h9.5a.5.5 0 0 0 .5-.5v-14a.5.5 0 0 0-.5-.5ZM16 5a1 1 0 0 1 1 1v4a1 1 0 0 1-1 1h-5a1 1 0 0 1-1-1V6a1 1 0 0 1 1-1h5Zm-1 2h-3v2h3V7Z"/></symbol><symbol id="icon-eds-i-mail-medium" viewBox="0 0 24 24"><path d="M20.462 3C21.875 3 23 4.184 23 5.619v12.762C23 19.816 21.875 21 20.462 21H3.538C2.125 21 1 19.816 1 18.381V5.619C1 4.184 2.125 3 3.538 3h16.924ZM21 8.158l-7.378 6.258a2.549 2.549 0 0 1-3.253-.008L3 8.16v10.222c0 .353.253.619.538.619h16.924c.285 0 .538-.266.538-.619V8.158ZM20.462 5H3.538c-.264 0-.5.228-.534.542l8.65 7.334c.2.165.492.165.684.007l8.656-7.342-.001-.025c-.044-.3-.274-.516-.531-.516Z"/></symbol><symbol id="icon-eds-i-mail-send-medium" viewBox="0 0 24 24"><path d="M20.444 5a2.562 2.562 0 0 1 2.548 2.37l.007.078.001.123v7.858A2.564 2.564 0 0 1 20.444 18H9.556A2.564 2.564 0 0 1 7 15.429l.001-7.977.007-.082A2.561 2.561 0 0 1 9.556 5h10.888ZM21 9.331l-5.46 3.51a1 1 0 0 1-1.08 0L9 9.332v6.097c0 .317.251.571.556.571h10.888a.564.564 0 0 0 .556-.571V9.33ZM20.444 7H9.556a.543.543 0 0 0-.32.105l5.763 3.706 5.766-3.706a.543.543 0 0 0-.32-.105ZM4.308 5a1 1 0 1 1 0 2H2a1 1 0 1 1 0-2h2.308Zm0 5.5a1 1 0 0 1 0 2H2a1 1 0 0 1 0-2h2.308Zm0 5.5a1 1 0 0 1 0 2H2a1 1 0 0 1 0-2h2.308Z"/></symbol><symbol id="icon-eds-i-mentions-medium" viewBox="0 0 24 24"><path d="m9.452 1.293 5.92 5.92 2.92-2.92a1 1 0 0 1 1.415 1.414l-2.92 2.92 5.92 5.92a1 1 0 0 1 0 1.415 10.371 10.371 0 0 1-10.378 2.584l.652 3.258A1 1 0 0 1 12 23H2a1 1 0 0 1-.874-1.486l4.789-8.62C4.194 9.074 4.9 4.43 8.038 1.292a1 1 0 0 1 1.414 0Zm-2.355 13.59L3.699 21h7.081l-.689-3.442a10.392 10.392 0 0 1-2.775-2.396l-.22-.28Zm1.69-11.427-.07.09a8.374 8.374 0 0 0 11.737 11.737l.089-.071L8.787 3.456Z"/></symbol><symbol id="icon-eds-i-menu-medium" viewBox="0 0 24 24"><path d="M21 4a1 1 0 0 1 0 2H3a1 1 0 1 1 0-2h18Zm-4 7a1 1 0 0 1 0 2H3a1 1 0 0 1 0-2h14Zm4 7a1 1 0 0 1 0 2H3a1 1 0 0 1 0-2h18Z"/></symbol><symbol id="icon-eds-i-metrics-medium" viewBox="0 0 24 24"><path d="M3 22a1 1 0 0 1-1-1V3a1 1 0 0 1 1-1h6a1 1 0 0 1 1 1v7h4V8a1 1 0 0 1 1-1h6a1 1 0 0 1 1 1v13a1 1 0 0 1-.883.993L21 22H3Zm17-2V9h-4v11h4Zm-6-8h-4v8h4v-8ZM8 4H4v16h4V4Z"/></symbol><symbol id="icon-eds-i-news-medium" viewBox="0 0 24 24"><path d="M17.384 3c.975 0 1.77.787 1.77 1.762v13.333c0 .462.354.846.815.899l.107.006.109-.006a.915.915 0 0 0 .809-.794l.006-.105V8.19a1 1 0 0 1 2 0v9.905A2.914 2.914 0 0 1 20.077 21H3.538a2.547 2.547 0 0 1-1.644-.601l-.147-.135A2.516 2.516 0 0 1 1 18.476V4.762C1 3.787 1.794 3 2.77 3h14.614Zm-.231 2H3v13.476c0 .11.035.216.1.304l.054.063c.101.1.24.157.384.157l13.761-.001-.026-.078a2.88 2.88 0 0 1-.115-.655l-.004-.17L17.153 5ZM14 15.021a.979.979 0 1 1 0 1.958H6a.979.979 0 1 1 0-1.958h8Zm0-8c.54 0 .979.438.979.979v4c0 .54-.438.979-.979.979H6A.979.979 0 0 1 5.021 12V8c0-.54.438-.979.979-.979h8Zm-.98 1.958H6.979v2.041h6.041V8.979Z"/></symbol><symbol id="icon-eds-i-newsletter-medium" viewBox="0 0 24 24"><path d="M21 10a1 1 0 0 1 1 1v9.5a2.5 2.5 0 0 1-2.5 2.5h-15A2.5 2.5 0 0 1 2 20.5V11a1 1 0 0 1 2 0v.439l8 4.888 8-4.889V11a1 1 0 0 1 1-1Zm-1 3.783-7.479 4.57a1 1 0 0 1-1.042 0l-7.48-4.57V20.5a.5.5 0 0 0 .501.5h15a.5.5 0 0 0 .5-.5v-6.717ZM15 9a1 1 0 0 1 0 2H9a1 1 0 0 1 0-2h6Zm2.5-8A2.5 2.5 0 0 1 20 3.5V9a1 1 0 0 1-2 0V3.5a.5.5 0 0 0-.5-.5h-11a.5.5 0 0 0-.5.5V9a1 1 0 1 1-2 0V3.5A2.5 2.5 0 0 1 6.5 1h11ZM15 5a1 1 0 0 1 0 2H9a1 1 0 1 1 0-2h6Z"/></symbol><symbol id="icon-eds-i-notifcation-medium" viewBox="0 0 24 24"><path d="M14 20a1 1 0 0 1 0 2h-4a1 1 0 0 1 0-2h4ZM3 18l-.133-.007c-1.156-.124-1.156-1.862 0-1.986l.3-.012C4.32 15.923 5 15.107 5 14V9.5C5 5.368 8.014 2 12 2s7 3.368 7 7.5V14c0 1.107.68 1.923 1.832 1.995l.301.012c1.156.124 1.156 1.862 0 1.986L21 18H3Zm9-14C9.17 4 7 6.426 7 9.5V14c0 .671-.146 1.303-.416 1.858L6.51 16h10.979l-.073-.142a4.192 4.192 0 0 1-.412-1.658L17 14V9.5C17 6.426 14.83 4 12 4Z"/></symbol><symbol id="icon-eds-i-publish-medium" viewBox="0 0 24 24"><g><path d="M16.296 1.291A1 1 0 0 0 15.591 1H5.545A2.542 2.542 0 0 0 3 3.538V13a1 1 0 1 0 2 0V3.538l.007-.087A.543.543 0 0 1 5.545 3h9.633L20 7.8v12.662a.534.534 0 0 1-.158.379.548.548 0 0 1-.387.159H11a1 1 0 1 0 0 2h8.455c.674 0 1.32-.267 1.798-.742A2.534 2.534 0 0 0 22 20.462V7.385a1 1 0 0 0-.294-.709l-5.41-5.385Z"/><path d="M10.762 16.647a1 1 0 0 0-1.525-1.294l-4.472 5.271-2.153-1.665a1 1 0 1 0-1.224 1.582l2.91 2.25a1 1 0 0 0 1.374-.144l5.09-6ZM16 10a1 1 0 1 1 0 2H8a1 1 0 1 1 0-2h8ZM12 7a1 1 0 0 0-1-1H8a1 1 0 1 0 0 2h3a1 1 0 0 0 1-1Z"/></g></symbol><symbol id="icon-eds-i-refresh-medium" viewBox="0 0 24 24"><g><path d="M7.831 5.636H6.032A8.76 8.76 0 0 1 9 3.631 8.549 8.549 0 0 1 12.232 3c.603 0 1.192.063 1.76.182C17.979 4.017 21 7.632 21 12a1 1 0 1 0 2 0c0-5.296-3.674-9.746-8.591-10.776A10.61 10.61 0 0 0 5 3.851V2.805a1 1 0 0 0-.987-1H4a1 1 0 0 0-1 1v3.831a1 1 0 0 0 1 1h3.831a1 1 0 0 0 .013-2h-.013ZM17.968 18.364c-1.59 1.632-3.784 2.636-6.2 2.636C6.948 21 3 16.993 3 12a1 1 0 1 0-2 0c0 6.053 4.799 11 10.768 11 2.788 0 5.324-1.082 7.232-2.85v1.045a1 1 0 1 0 2 0v-3.831a1 1 0 0 0-1-1h-3.831a1 1 0 0 0 0 2h1.799Z"/></g></symbol><symbol id="icon-eds-i-search-medium" viewBox="0 0 24 24"><path d="M11 1c5.523 0 10 4.477 10 10 0 2.4-.846 4.604-2.256 6.328l3.963 3.965a1 1 0 0 1-1.414 1.414l-3.965-3.963A9.959 9.959 0 0 1 11 21C5.477 21 1 16.523 1 11S5.477 1 11 1Zm0 2a8 8 0 1 0 0 16 8 8 0 0 0 0-16Z"/></symbol><symbol id="icon-eds-i-settings-medium" viewBox="0 0 24 24"><path d="M11.382 1h1.24a2.508 2.508 0 0 1 2.334 1.63l.523 1.378 1.59.933 1.444-.224c.954-.132 1.89.3 2.422 1.101l.095.155.598 1.066a2.56 2.56 0 0 1-.195 2.848l-.894 1.161v1.896l.92 1.163c.6.768.707 1.812.295 2.674l-.09.17-.606 1.08a2.504 2.504 0 0 1-2.531 1.25l-1.428-.223-1.589.932-.523 1.378a2.512 2.512 0 0 1-2.155 1.625L12.65 23h-1.27a2.508 2.508 0 0 1-2.334-1.63l-.524-1.379-1.59-.933-1.443.225c-.954.132-1.89-.3-2.422-1.101l-.095-.155-.598-1.066a2.56 2.56 0 0 1 .195-2.847l.891-1.161v-1.898l-.919-1.162a2.562 2.562 0 0 1-.295-2.674l.09-.17.606-1.08a2.504 2.504 0 0 1 2.531-1.25l1.43.223 1.618-.938.524-1.375.07-.167A2.507 2.507 0 0 1 11.382 1Zm.003 2a.509.509 0 0 0-.47.338l-.65 1.71a1 1 0 0 1-.434.51L7.6 6.85a1 1 0 0 1-.655.123l-1.762-.275a.497.497 0 0 0-.498.252l-.61 1.088a.562.562 0 0 0 .04.619l1.13 1.43a1 1 0 0 1 .216.62v2.585a1 1 0 0 1-.207.61L4.15 15.339a.568.568 0 0 0-.036.634l.601 1.072a.494.494 0 0 0 .484.26l1.78-.278a1 1 0 0 1 .66.126l2.2 1.292a1 1 0 0 1 .43.507l.648 1.71a.508.508 0 0 0 .467.338h1.263a.51.51 0 0 0 .47-.34l.65-1.708a1 1 0 0 1 .428-.507l2.201-1.292a1 1 0 0 1 .66-.126l1.763.275a.497.497 0 0 0 .498-.252l.61-1.088a.562.562 0 0 0-.04-.619l-1.13-1.43a1 1 0 0 1-.216-.62v-2.585a1 1 0 0 1 .207-.61l1.105-1.437a.568.568 0 0 0 .037-.634l-.601-1.072a.494.494 0 0 0-.484-.26l-1.78.278a1 1 0 0 1-.66-.126l-2.2-1.292a1 1 0 0 1-.43-.507l-.649-1.71A.508.508 0 0 0 12.62 3h-1.234ZM12 8a4 4 0 1 1 0 8 4 4 0 0 1 0-8Zm0 2a2 2 0 1 0 0 4 2 2 0 0 0 0-4Z"/></symbol><symbol id="icon-eds-i-shipping-medium" viewBox="0 0 24 24"><path d="M16.515 2c1.406 0 2.706.728 3.352 1.902l2.02 3.635.02.042.036.089.031.105.012.058.01.073.004.075v11.577c0 .64-.244 1.255-.683 1.713a2.356 2.356 0 0 1-1.701.731H4.386a2.356 2.356 0 0 1-1.702-.731 2.476 2.476 0 0 1-.683-1.713V7.948c.01-.217.083-.43.22-.6L4.2 3.905C4.833 2.755 6.089 2.032 7.486 2h9.029ZM20 9H4v10.556a.49.49 0 0 0 .075.26l.053.07a.356.356 0 0 0 .257.114h15.23c.094 0 .186-.04.258-.115a.477.477 0 0 0 .127-.33V9Zm-2 7.5a1 1 0 0 1 0 2h-4a1 1 0 0 1 0-2h4ZM16.514 4H13v3h6.3l-1.183-2.13c-.288-.522-.908-.87-1.603-.87ZM11 3.999H7.51c-.679.017-1.277.36-1.566.887L4.728 7H11V3.999Z"/></symbol><symbol id="icon-eds-i-step-guide-medium" viewBox="0 0 24 24"><path d="M11.394 9.447a1 1 0 1 0-1.788-.894l-.88 1.759-.019-.02a1 1 0 1 0-1.414 1.415l1 1a1 1 0 0 0 1.601-.26l1.5-3ZM12 11a1 1 0 0 1 1-1h3a1 1 0 1 1 0 2h-3a1 1 0 0 1-1-1ZM12 17a1 1 0 0 1 1-1h3a1 1 0 1 1 0 2h-3a1 1 0 0 1-1-1ZM10.947 14.105a1 1 0 0 1 .447 1.342l-1.5 3a1 1 0 0 1-1.601.26l-1-1a1 1 0 1 1 1.414-1.414l.02.019.879-1.76a1 1 0 0 1 1.341-.447Z"/><path d="M5.545 1A2.542 2.542 0 0 0 3 3.538v16.924A2.542 2.542 0 0 0 5.545 23h12.91A2.542 2.542 0 0 0 21 20.462V7.5a1 1 0 0 0-.293-.707l-5.5-5.5A1 1 0 0 0 14.5 1H5.545ZM5 3.538C5 3.245 5.24 3 5.545 3h8.54L19 7.914v12.547c0 .294-.24.539-.546.539H5.545A.542.542 0 0 1 5 20.462V3.538Z" clip-rule="evenodd"/></symbol><symbol id="icon-eds-i-submission-medium" viewBox="0 0 24 24"><g><path d="M5 3.538C5 3.245 5.24 3 5.545 3h9.633L20 7.8v12.662a.535.535 0 0 1-.158.379.549.549 0 0 1-.387.159H6a1 1 0 0 1-1-1v-2.5a1 1 0 1 0-2 0V20a3 3 0 0 0 3 3h13.455c.673 0 1.32-.266 1.798-.742A2.535 2.535 0 0 0 22 20.462V7.385a1 1 0 0 0-.294-.709l-5.41-5.385A1 1 0 0 0 15.591 1H5.545A2.542 2.542 0 0 0 3 3.538V7a1 1 0 0 0 2 0V3.538Z"/><path d="m13.707 13.707-4 4a1 1 0 0 1-1.414 0l-.083-.094a1 1 0 0 1 .083-1.32L10.585 14 2 14a1 1 0 1 1 0-2l8.583.001-2.29-2.294a1 1 0 0 1 1.414-1.414l4.037 4.04.043.05.043.06.059.098.03.063.031.085.03.113.017.122L14 13l-.004.087-.017.118-.013.056-.034.104-.049.105-.048.081-.07.093-.058.063Z"/></g></symbol><symbol id="icon-eds-i-table-1-medium" viewBox="0 0 24 24"><path d="M4.385 22a2.56 2.56 0 0 1-1.14-.279C2.485 21.341 2 20.614 2 19.615V4.385c0-.315.067-.716.279-1.14C2.659 2.485 3.386 2 4.385 2h15.23c.315 0 .716.067 1.14.279.76.38 1.245 1.107 1.245 2.106v15.23c0 .315-.067.716-.279 1.14-.38.76-1.107 1.245-2.106 1.245H4.385ZM4 19.615c0 .213.034.265.14.317a.71.71 0 0 0 .245.068H8v-4H4v3.615ZM20 16H10v4h9.615c.213 0 .265-.034.317-.14a.71.71 0 0 0 .068-.245V16Zm0-2v-4H10v4h10ZM4 14h4v-4H4v4ZM19.615 4H10v4h10V4.385c0-.213-.034-.265-.14-.317A.71.71 0 0 0 19.615 4ZM8 4H4.385l-.082.002c-.146.01-.19.047-.235.138A.71.71 0 0 0 4 4.385V8h4V4Z"/></symbol><symbol id="icon-eds-i-table-2-medium" viewBox="0 0 24 24"><path d="M4.384 22A2.384 2.384 0 0 1 2 19.616V4.384A2.384 2.384 0 0 1 4.384 2h15.232A2.384 2.384 0 0 1 22 4.384v15.232A2.384 2.384 0 0 1 19.616 22H4.384ZM10 15H4v4.616c0 .212.172.384.384.384H10v-5Zm5 0h-3v5h3v-5Zm5 0h-3v5h2.616a.384.384 0 0 0 .384-.384V15ZM10 9H4v4h6V9Zm5 0h-3v4h3V9Zm5 0h-3v4h3V9Zm-.384-5H4.384A.384.384 0 0 0 4 4.384V7h16V4.384A.384.384 0 0 0 19.616 4Z"/></symbol><symbol id="icon-eds-i-tag-medium" viewBox="0 0 24 24"><path d="m12.621 1.998.127.004L20.496 2a1.5 1.5 0 0 1 1.497 1.355L22 3.5l-.005 7.669c.038.456-.133.905-.447 1.206l-9.02 9.018a2.075 2.075 0 0 1-2.932 0l-6.99-6.99a2.075 2.075 0 0 1 .001-2.933L11.61 2.47c.246-.258.573-.418.881-.46l.131-.011Zm.286 2-8.885 8.886a.075.075 0 0 0 0 .106l6.987 6.988c.03.03.077.03.106 0l8.883-8.883L19.999 4l-7.092-.002ZM16 6.5a1.5 1.5 0 0 1 .144 2.993L16 9.5a1.5 1.5 0 0 1 0-3Z"/></symbol><symbol id="icon-eds-i-trash-medium" viewBox="0 0 24 24"><path d="M12 1c2.717 0 4.913 2.232 4.997 5H21a1 1 0 0 1 0 2h-1v12.5c0 1.389-1.152 2.5-2.556 2.5H6.556C5.152 23 4 21.889 4 20.5V8H3a1 1 0 1 1 0-2h4.003l.001-.051C7.114 3.205 9.3 1 12 1Zm6 7H6v12.5c0 .238.19.448.454.492l.102.008h10.888c.315 0 .556-.232.556-.5V8Zm-4 3a1 1 0 0 1 1 1v6.005a1 1 0 0 1-2 0V12a1 1 0 0 1 1-1Zm-4 0a1 1 0 0 1 1 1v6a1 1 0 0 1-2 0v-6a1 1 0 0 1 1-1Zm2-8c-1.595 0-2.914 1.32-2.996 3h5.991v-.02C14.903 4.31 13.589 3 12 3Z"/></symbol><symbol id="icon-eds-i-user-account-medium" viewBox="0 0 24 24"><path d="M12 1c6.075 0 11 4.925 11 11s-4.925 11-11 11S1 18.075 1 12 5.925 1 12 1Zm0 16c-1.806 0-3.52.994-4.664 2.698A8.947 8.947 0 0 0 12 21a8.958 8.958 0 0 0 4.664-1.301C15.52 17.994 13.806 17 12 17Zm0-14a9 9 0 0 0-6.25 15.476C7.253 16.304 9.54 15 12 15s4.747 1.304 6.25 3.475A9 9 0 0 0 12 3Zm0 3a4 4 0 1 1 0 8 4 4 0 0 1 0-8Zm0 2a2 2 0 1 0 0 4 2 2 0 0 0 0-4Z"/></symbol><symbol id="icon-eds-i-user-add-medium" viewBox="0 0 24 24"><path d="M9 1a5 5 0 1 1 0 10A5 5 0 0 1 9 1Zm0 2a3 3 0 1 0 0 6 3 3 0 0 0 0-6Zm9 10a1 1 0 0 1 1 1v3h3a1 1 0 0 1 0 2h-3v3a1 1 0 0 1-2 0v-3h-3a1 1 0 0 1 0-2h3v-3a1 1 0 0 1 1-1Zm-5.545-.15a1 1 0 1 1-.91 1.78 5.713 5.713 0 0 0-5.705.282c-1.67 1.068-2.728 2.927-2.832 4.956L3.004 20 11.5 20a1 1 0 0 1 .993.883L12.5 21a1 1 0 0 1-1 1H2a1 1 0 0 1-1-1v-.876c.028-2.812 1.446-5.416 3.763-6.897a7.713 7.713 0 0 1 7.692-.378Z"/></symbol><symbol id="icon-eds-i-user-assign-medium" viewBox="0 0 24 24"><path d="M16.226 13.298a1 1 0 0 1 1.414-.01l.084.093a1 1 0 0 1-.073 1.32L15.39 17H22a1 1 0 0 1 0 2h-6.611l2.262 2.298a1 1 0 0 1-1.425 1.404l-3.939-4a1 1 0 0 1 0-1.404l3.94-4Zm-3.771-.449a1 1 0 1 1-.91 1.781 5.713 5.713 0 0 0-5.705.282c-1.67 1.068-2.728 2.927-2.832 4.956L3.004 20 10.5 20a1 1 0 0 1 .993.883L11.5 21a1 1 0 0 1-1 1H2a1 1 0 0 1-1-1v-.876c.028-2.812 1.446-5.416 3.763-6.897a7.713 7.713 0 0 1 7.692-.378ZM9 1a5 5 0 1 1 0 10A5 5 0 0 1 9 1Zm0 2a3 3 0 1 0 0 6 3 3 0 0 0 0-6Z"/></symbol><symbol id="icon-eds-i-user-block-medium" viewBox="0 0 24 24"><path d="M9 1a5 5 0 1 1 0 10A5 5 0 0 1 9 1Zm0 2a3 3 0 1 0 0 6 3 3 0 0 0 0-6Zm9 10a5 5 0 1 1 0 10 5 5 0 0 1 0-10Zm-5.545-.15a1 1 0 1 1-.91 1.78 5.713 5.713 0 0 0-5.705.282c-1.67 1.068-2.728 2.927-2.832 4.956L3.004 20 11.5 20a1 1 0 0 1 .993.883L12.5 21a1 1 0 0 1-1 1H2a1 1 0 0 1-1-1v-.876c.028-2.812 1.446-5.416 3.763-6.897a7.713 7.713 0 0 1 7.692-.378ZM15 18a3 3 0 0 0 4.294 2.707l-4.001-4c-.188.391-.293.83-.293 1.293Zm3-3c-.463 0-.902.105-1.294.293l4.001 4A3 3 0 0 0 18 15Z"/></symbol><symbol id="icon-eds-i-user-check-medium" viewBox="0 0 24 24"><path d="M9 1a5 5 0 1 1 0 10A5 5 0 0 1 9 1Zm0 2a3 3 0 1 0 0 6 3 3 0 0 0 0-6Zm13.647 12.237a1 1 0 0 1 .116 1.41l-5.091 6a1 1 0 0 1-1.375.144l-2.909-2.25a1 1 0 1 1 1.224-1.582l2.153 1.665 4.472-5.271a1 1 0 0 1 1.41-.116Zm-8.139-.977c.22.214.428.44.622.678a1 1 0 1 1-1.548 1.266 6.025 6.025 0 0 0-1.795-1.49.86.86 0 0 1-.163-.048l-.079-.036a5.721 5.721 0 0 0-2.62-.63l-.194.006c-2.76.134-5.022 2.177-5.592 4.864l-.035.175-.035.213c-.03.201-.05.405-.06.61L3.003 20 10 20a1 1 0 0 1 .993.883L11 21a1 1 0 0 1-1 1H2a1 1 0 0 1-1-1v-.876l.005-.223.02-.356.02-.222.03-.248.022-.15c.02-.133.044-.265.071-.397.44-2.178 1.725-4.105 3.595-5.301a7.75 7.75 0 0 1 3.755-1.215l.12-.004a7.908 7.908 0 0 1 5.87 2.252Z"/></symbol><symbol id="icon-eds-i-user-delete-medium" viewBox="0 0 24 24"><path d="M9 1a5 5 0 1 1 0 10A5 5 0 0 1 9 1Zm0 2a3 3 0 1 0 0 6 3 3 0 0 0 0-6ZM4.763 13.227a7.713 7.713 0 0 1 7.692-.378 1 1 0 1 1-.91 1.781 5.713 5.713 0 0 0-5.705.282c-1.67 1.068-2.728 2.927-2.832 4.956L3.004 20H11.5a1 1 0 0 1 .993.883L12.5 21a1 1 0 0 1-1 1H2a1 1 0 0 1-1-1v-.876c.028-2.812 1.446-5.416 3.763-6.897Zm11.421 1.543 2.554 2.553 2.555-2.553a1 1 0 0 1 1.414 1.414l-2.554 2.554 2.554 2.555a1 1 0 0 1-1.414 1.414l-2.555-2.554-2.554 2.554a1 1 0 0 1-1.414-1.414l2.553-2.555-2.553-2.554a1 1 0 0 1 1.414-1.414Z"/></symbol><symbol id="icon-eds-i-user-edit-medium" viewBox="0 0 24 24"><path d="m19.876 10.77 2.831 2.83a1 1 0 0 1 0 1.415l-7.246 7.246a1 1 0 0 1-.572.284l-3.277.446a1 1 0 0 1-1.125-1.13l.461-3.277a1 1 0 0 1 .283-.567l7.23-7.246a1 1 0 0 1 1.415-.001Zm-7.421 2.08a1 1 0 1 1-.91 1.78 5.713 5.713 0 0 0-5.705.282c-1.67 1.068-2.728 2.927-2.832 4.956L3.004 20 7.5 20a1 1 0 0 1 .993.883L8.5 21a1 1 0 0 1-1 1H2a1 1 0 0 1-1-1v-.876c.028-2.812 1.446-5.416 3.763-6.897a7.713 7.713 0 0 1 7.692-.378Zm6.715.042-6.29 6.3-.23 1.639 1.633-.222 6.302-6.302-1.415-1.415ZM9 1a5 5 0 1 1 0 10A5 5 0 0 1 9 1Zm0 2a3 3 0 1 0 0 6 3 3 0 0 0 0-6Z"/></symbol><symbol id="icon-eds-i-user-linked-medium" viewBox="0 0 24 24"><path d="M15.65 6c.31 0 .706.066 1.122.274C17.522 6.65 18 7.366 18 8.35v12.3c0 .31-.066.706-.274 1.122-.375.75-1.092 1.228-2.076 1.228H3.35a2.52 2.52 0 0 1-1.122-.274C1.478 22.35 1 21.634 1 20.65V8.35c0-.31.066-.706.274-1.122C1.65 6.478 2.366 6 3.35 6h12.3Zm0 2-12.376.002c-.134.007-.17.04-.21.12A.672.672 0 0 0 3 8.35v12.3c0 .198.028.24.122.287.09.044.2.063.228.063h.887c.788-2.269 2.814-3.5 5.263-3.5 2.45 0 4.475 1.231 5.263 3.5h.887c.198 0 .24-.028.287-.122.044-.09.063-.2.063-.228V8.35c0-.198-.028-.24-.122-.287A.672.672 0 0 0 15.65 8ZM9.5 19.5c-1.36 0-2.447.51-3.06 1.5h6.12c-.613-.99-1.7-1.5-3.06-1.5ZM20.65 1A2.35 2.35 0 0 1 23 3.348V15.65A2.35 2.35 0 0 1 20.65 18H20a1 1 0 0 1 0-2h.65a.35.35 0 0 0 .35-.35V3.348A.35.35 0 0 0 20.65 3H8.35a.35.35 0 0 0-.35.348V4a1 1 0 1 1-2 0v-.652A2.35 2.35 0 0 1 8.35 1h12.3ZM9.5 10a3.5 3.5 0 1 1 0 7 3.5 3.5 0 0 1 0-7Zm0 2a1.5 1.5 0 1 0 0 3 1.5 1.5 0 0 0 0-3Z"/></symbol><symbol id="icon-eds-i-user-multiple-medium" viewBox="0 0 24 24"><path d="M9 1a5 5 0 1 1 0 10A5 5 0 0 1 9 1Zm6 0a5 5 0 0 1 0 10 1 1 0 0 1-.117-1.993L15 9a3 3 0 0 0 0-6 1 1 0 0 1 0-2ZM9 3a3 3 0 1 0 0 6 3 3 0 0 0 0-6Zm8.857 9.545a7.99 7.99 0 0 1 2.651 1.715A8.31 8.31 0 0 1 23 20.134V21a1 1 0 0 1-1 1h-3a1 1 0 0 1 0-2h1.995l-.005-.153a6.307 6.307 0 0 0-1.673-3.945l-.204-.209a5.99 5.99 0 0 0-1.988-1.287 1 1 0 1 1 .732-1.861Zm-3.349 1.715A8.31 8.31 0 0 1 17 20.134V21a1 1 0 0 1-1 1H2a1 1 0 0 1-1-1v-.877c.044-4.343 3.387-7.908 7.638-8.115a7.908 7.908 0 0 1 5.87 2.252ZM9.016 14l-.285.006c-3.104.15-5.58 2.718-5.725 5.9L3.004 20h11.991l-.005-.153a6.307 6.307 0 0 0-1.673-3.945l-.204-.209A5.924 5.924 0 0 0 9.3 14.008L9.016 14Z"/></symbol><symbol id="icon-eds-i-user-notify-medium" viewBox="0 0 24 24"><path d="M9 1a5 5 0 1 1 0 10A5 5 0 0 1 9 1Zm0 2a3 3 0 1 0 0 6 3 3 0 0 0 0-6Zm10 18v1a1 1 0 0 1-2 0v-1h-3a1 1 0 0 1 0-2v-2.818C14 13.885 15.777 12 18 12s4 1.885 4 4.182V19a1 1 0 0 1 0 2h-3Zm-6.545-8.15a1 1 0 1 1-.91 1.78 5.713 5.713 0 0 0-5.705.282c-1.67 1.068-2.728 2.927-2.832 4.956L3.004 20 11.5 20a1 1 0 0 1 .993.883L12.5 21a1 1 0 0 1-1 1H2a1 1 0 0 1-1-1v-.876c.028-2.812 1.446-5.416 3.763-6.897a7.713 7.713 0 0 1 7.692-.378ZM18 14c-1.091 0-2 .964-2 2.182V19h4v-2.818c0-1.165-.832-2.098-1.859-2.177L18 14Z"/></symbol><symbol id="icon-eds-i-user-remove-medium" viewBox="0 0 24 24"><path d="M9 1a5 5 0 1 1 0 10A5 5 0 0 1 9 1Zm0 2a3 3 0 1 0 0 6 3 3 0 0 0 0-6Zm3.455 9.85a1 1 0 1 1-.91 1.78 5.713 5.713 0 0 0-5.705.282c-1.67 1.068-2.728 2.927-2.832 4.956L3.004 20 11.5 20a1 1 0 0 1 .993.883L12.5 21a1 1 0 0 1-1 1H2a1 1 0 0 1-1-1v-.876c.028-2.812 1.446-5.416 3.763-6.897a7.713 7.713 0 0 1 7.692-.378ZM22 17a1 1 0 0 1 0 2h-8a1 1 0 0 1 0-2h8Z"/></symbol><symbol id="icon-eds-i-user-single-medium" viewBox="0 0 24 24"><path d="M12 1a5 5 0 1 1 0 10 5 5 0 0 1 0-10Zm0 2a3 3 0 1 0 0 6 3 3 0 0 0 0-6Zm-.406 9.008a8.965 8.965 0 0 1 6.596 2.494A9.161 9.161 0 0 1 21 21.025V22a1 1 0 0 1-1 1H4a1 1 0 0 1-1-1v-.985c.05-4.825 3.815-8.777 8.594-9.007Zm.39 1.992-.299.006c-3.63.175-6.518 3.127-6.678 6.775L5 21h13.998l-.009-.268a7.157 7.157 0 0 0-1.97-4.573l-.214-.213A6.967 6.967 0 0 0 11.984 14Z"/></symbol><symbol id="icon-eds-i-warning-circle-medium" viewBox="0 0 24 24"><path d="M12 1c6.075 0 11 4.925 11 11s-4.925 11-11 11S1 18.075 1 12 5.925 1 12 1Zm0 2a9 9 0 1 0 0 18 9 9 0 0 0 0-18Zm0 11.5a1.5 1.5 0 0 1 .144 2.993L12 17.5a1.5 1.5 0 0 1 0-3ZM12 6a1 1 0 0 1 1 1v5a1 1 0 0 1-2 0V7a1 1 0 0 1 1-1Z"/></symbol><symbol id="icon-eds-i-warning-filled-medium" viewBox="0 0 24 24"><path d="M12 1c6.075 0 11 4.925 11 11s-4.925 11-11 11S1 18.075 1 12 5.925 1 12 1Zm0 13.5a1.5 1.5 0 0 0 0 3l.144-.007A1.5 1.5 0 0 0 12 14.5ZM12 6a1 1 0 0 0-1 1v5a1 1 0 0 0 2 0V7a1 1 0 0 0-1-1Z"/></symbol><symbol id="icon-chevron-left-medium" viewBox="0 0 24 24"><path d="M15.7194 3.3054C15.3358 2.90809 14.7027 2.89699 14.3054 3.28061L6.54342 10.7757C6.19804 11.09 6 11.5335 6 12C6 12.4665 6.19804 12.91 6.5218 13.204L14.3054 20.7194C14.7027 21.103 15.3358 21.0919 15.7194 20.6946C16.103 20.2973 16.0919 19.6642 15.6946 19.2806L8.155 12L15.6946 4.71939C16.0614 4.36528 16.099 3.79863 15.8009 3.40105L15.7194 3.3054Z"/></symbol><symbol id="icon-chevron-right-medium" viewBox="0 0 24 24"><path d="M8.28061 3.3054C8.66423 2.90809 9.29729 2.89699 9.6946 3.28061L17.4566 10.7757C17.802 11.09 18 11.5335 18 12C18 12.4665 17.802 12.91 17.4782 13.204L9.6946 20.7194C9.29729 21.103 8.66423 21.0919 8.28061 20.6946C7.89699 20.2973 7.90809 19.6642 8.3054 19.2806L15.845 12L8.3054 4.71939C7.93865 4.36528 7.90098 3.79863 8.19908 3.40105L8.28061 3.3054Z"/></symbol><symbol id="icon-eds-alerts" viewBox="0 0 32 32"><path d="M28 12.667c.736 0 1.333.597 1.333 1.333v13.333A3.333 3.333 0 0 1 26 30.667H6a3.333 3.333 0 0 1-3.333-3.334V14a1.333 1.333 0 1 1 2.666 0v1.252L16 21.769l10.667-6.518V14c0-.736.597-1.333 1.333-1.333Zm-1.333 5.71-9.972 6.094c-.427.26-.963.26-1.39 0l-9.972-6.094v8.956c0 .368.299.667.667.667h20a.667.667 0 0 0 .667-.667v-8.956ZM19.333 12a1.333 1.333 0 1 1 0 2.667h-6.666a1.333 1.333 0 1 1 0-2.667h6.666Zm4-10.667a3.333 3.333 0 0 1 3.334 3.334v6.666a1.333 1.333 0 1 1-2.667 0V4.667A.667.667 0 0 0 23.333 4H8.667A.667.667 0 0 0 8 4.667v6.666a1.333 1.333 0 1 1-2.667 0V4.667a3.333 3.333 0 0 1 3.334-3.334h14.666Zm-4 5.334a1.333 1.333 0 0 1 0 2.666h-6.666a1.333 1.333 0 1 1 0-2.666h6.666Z"/></symbol><symbol id="icon-eds-arrow-up" viewBox="0 0 24 24"><path fill-rule="evenodd" d="m13.002 7.408 4.88 4.88a.99.99 0 0 0 1.32.08l.09-.08c.39-.39.39-1.03 0-1.42l-6.58-6.58a1.01 1.01 0 0 0-1.42 0l-6.58 6.58a1 1 0 0 0-.09 1.32l.08.1a1 1 0 0 0 1.42-.01l4.88-4.87v11.59a.99.99 0 0 0 .88.99l.12.01c.55 0 1-.45 1-1V7.408z" class="layer"/></symbol><symbol id="icon-eds-checklist" viewBox="0 0 32 32"><path d="M19.2 1.333a3.468 3.468 0 0 1 3.381 2.699L24.667 4C26.515 4 28 5.52 28 7.38v19.906c0 1.86-1.485 3.38-3.333 3.38H7.333c-1.848 0-3.333-1.52-3.333-3.38V7.38C4 5.52 5.485 4 7.333 4h2.093A3.468 3.468 0 0 1 12.8 1.333h6.4ZM9.426 6.667H7.333c-.36 0-.666.312-.666.713v19.906c0 .401.305.714.666.714h17.334c.36 0 .666-.313.666-.714V7.38c0-.4-.305-.713-.646-.714l-2.121.033A3.468 3.468 0 0 1 19.2 9.333h-6.4a3.468 3.468 0 0 1-3.374-2.666Zm12.715 5.606c.586.446.7 1.283.253 1.868l-7.111 9.334a1.333 1.333 0 0 1-1.792.306l-3.556-2.333a1.333 1.333 0 1 1 1.463-2.23l2.517 1.651 6.358-8.344a1.333 1.333 0 0 1 1.868-.252ZM19.2 4h-6.4a.8.8 0 0 0-.8.8v1.067a.8.8 0 0 0 .8.8h6.4a.8.8 0 0 0 .8-.8V4.8a.8.8 0 0 0-.8-.8Z"/></symbol><symbol id="icon-eds-citation" viewBox="0 0 36 36"><path d="M23.25 1.5a1.5 1.5 0 0 1 1.06.44l8.25 8.25a1.5 1.5 0 0 1 .44 1.06v19.5c0 2.105-1.645 3.75-3.75 3.75H18a1.5 1.5 0 0 1 0-3h11.25c.448 0 .75-.302.75-.75V11.873L22.628 4.5H8.31a.811.811 0 0 0-.8.68l-.011.13V16.5a1.5 1.5 0 0 1-3 0V5.31A3.81 3.81 0 0 1 8.31 1.5h14.94ZM8.223 20.358a.984.984 0 0 1-.192 1.378l-.048.034c-.54.36-.942.676-1.206.951-.59.614-.885 1.395-.885 2.343.115-.028.288-.042.518-.042.662 0 1.26.237 1.791.711.533.474.799 1.074.799 1.799 0 .753-.259 1.352-.777 1.799-.518.446-1.151.669-1.9.669-1.006 0-1.812-.293-2.417-.878C3.302 28.536 3 27.657 3 26.486c0-1.115.165-2.085.496-2.907.331-.823.734-1.513 1.209-2.071.475-.558.971-.997 1.49-1.318a6.01 6.01 0 0 1 .347-.2 1.321 1.321 0 0 1 1.681.368Zm7.5 0a.984.984 0 0 1-.192 1.378l-.048.034c-.54.36-.942.676-1.206.951-.59.614-.885 1.395-.885 2.343.115-.028.288-.042.518-.042.662 0 1.26.237 1.791.711.533.474.799 1.074.799 1.799 0 .753-.259 1.352-.777 1.799-.518.446-1.151.669-1.9.669-1.006 0-1.812-.293-2.417-.878-.604-.586-.906-1.465-.906-2.636 0-1.115.165-2.085.496-2.907.331-.823.734-1.513 1.209-2.071.475-.558.971-.997 1.49-1.318a6.01 6.01 0 0 1 .347-.2 1.321 1.321 0 0 1 1.681.368Z"/></symbol><symbol id="icon-eds-i-access-indicator" viewBox="0 0 16 16"><circle cx="4.5" cy="11.5" r="3.5" style="fill:currentColor"/><path fill-rule="evenodd" d="M4 3v3a1 1 0 0 1-2 0V2.923C2 1.875 2.84 1 3.909 1h5.909a1 1 0 0 1 .713.298l3.181 3.231a1 1 0 0 1 .288.702v7.846c0 .505-.197.993-.554 1.354a1.902 1.902 0 0 1-1.355.569H10a1 1 0 1 1 0-2h2V5.64L9.4 3H4Z" clip-rule="evenodd" style="fill:#222"/></symbol><symbol id="icon-eds-i-github-medium" viewBox="0 0 24 24"><path d="M 11.964844 0 C 5.347656 0 0 5.269531 0 11.792969 C 0 17.003906 3.425781 21.417969 8.179688 22.976562 C 8.773438 23.09375 8.992188 22.722656 8.992188 22.410156 C 8.992188 22.136719 8.972656 21.203125 8.972656 20.226562 C 5.644531 20.929688 4.953125 18.820312 4.953125 18.820312 C 4.417969 17.453125 3.625 17.101562 3.625 17.101562 C 2.535156 16.378906 3.703125 16.378906 3.703125 16.378906 C 4.914062 16.457031 5.546875 17.589844 5.546875 17.589844 C 6.617188 19.386719 8.339844 18.878906 9.03125 18.566406 C 9.132812 17.804688 9.449219 17.277344 9.785156 16.984375 C 7.132812 16.710938 4.339844 15.695312 4.339844 11.167969 C 4.339844 9.878906 4.8125 8.824219 5.566406 8.003906 C 5.445312 7.710938 5.03125 6.5 5.683594 4.878906 C 5.683594 4.878906 6.695312 4.566406 8.972656 6.089844 C 9.949219 5.832031 10.953125 5.703125 11.964844 5.699219 C 12.972656 5.699219 14.003906 5.835938 14.957031 6.089844 C 17.234375 4.566406 18.242188 4.878906 18.242188 4.878906 C 18.898438 6.5 18.480469 7.710938 18.363281 8.003906 C 19.136719 8.824219 19.589844 9.878906 19.589844 11.167969 C 19.589844 15.695312 16.796875 16.691406 14.125 16.984375 C 14.558594 17.355469 14.933594 18.058594 14.933594 19.171875 C 14.933594 20.753906 14.914062 22.019531 14.914062 22.410156 C 14.914062 22.722656 15.132812 23.09375 15.726562 22.976562 C 20.480469 21.414062 23.910156 17.003906 23.910156 11.792969 C 23.929688 5.269531 18.558594 0 11.964844 0 Z M 11.964844 0 "/></symbol><symbol id="icon-eds-i-limited-access" viewBox="0 0 16 16"><path fill-rule="evenodd" d="M4 3v3a1 1 0 0 1-2 0V2.923C2 1.875 2.84 1 3.909 1h5.909a1 1 0 0 1 .713.298l3.181 3.231a1 1 0 0 1 .288.702V6a1 1 0 1 1-2 0v-.36L9.4 3H4ZM3 8a1 1 0 0 1 1 1v1a1 1 0 1 1-2 0V9a1 1 0 0 1 1-1Zm10 0a1 1 0 0 1 1 1v1a1 1 0 1 1-2 0V9a1 1 0 0 1 1-1Zm-3.5 6a1 1 0 0 1-1 1h-1a1 1 0 1 1 0-2h1a1 1 0 0 1 1 1Zm2.441-1a1 1 0 0 1 2 0c0 .73-.246 1.306-.706 1.664a1.61 1.61 0 0 1-.876.334l-.032.002H11.5a1 1 0 1 1 0-2h.441ZM4 13a1 1 0 0 0-2 0c0 .73.247 1.306.706 1.664a1.609 1.609 0 0 0 .876.334l.032.002H4.5a1 1 0 1 0 0-2H4Z" clip-rule="evenodd"/></symbol><symbol id="icon-eds-i-subjects-medium" viewBox="0 0 24 24"><g id="icon-subjects-copy" stroke="none" stroke-width="1" fill-rule="evenodd"><path d="M13.3846154,2 C14.7015971,2 15.7692308,3.06762994 15.7692308,4.38461538 L15.7692308,7.15384615 C15.7692308,8.47082629 14.7015955,9.53846154 13.3846154,9.53846154 L13.1038388,9.53925278 C13.2061091,9.85347965 13.3815528,10.1423885 13.6195822,10.3804178 C13.9722182,10.7330539 14.436524,10.9483278 14.9293854,10.9918129 L15.1153846,11 C16.2068332,11 17.2535347,11.433562 18.0254647,12.2054189 C18.6411944,12.8212361 19.0416785,13.6120766 19.1784166,14.4609738 L19.6153846,14.4615385 C20.932386,14.4615385 22,15.5291672 22,16.8461538 L22,19.6153846 C22,20.9323924 20.9323924,22 19.6153846,22 L16.8461538,22 C15.5291672,22 14.4615385,20.932386 14.4615385,19.6153846 L14.4615385,16.8461538 C14.4615385,15.5291737 15.5291737,14.4615385 16.8461538,14.4615385 L17.126925,14.460779 C17.0246537,14.1465537 16.8492179,13.857633 16.6112344,13.6196157 C16.2144418,13.2228606 15.6764136,13 15.1153846,13 C14.0239122,13 12.9771569,12.5664197 12.2053686,11.7946314 C12.1335167,11.7227795 12.0645962,11.6485444 11.9986839,11.5721119 C11.9354038,11.6485444 11.8664833,11.7227795 11.7946314,11.7946314 C11.0228431,12.5664197 9.97608778,13 8.88461538,13 C8.323576,13 7.78552852,13.2228666 7.38881294,13.6195822 C7.15078359,13.8576115 6.97533988,14.1465203 6.8730696,14.4607472 L7.15384615,14.4615385 C8.47082629,14.4615385 9.53846154,15.5291737 9.53846154,16.8461538 L9.53846154,19.6153846 C9.53846154,20.932386 8.47083276,22 7.15384615,22 L4.38461538,22 C3.06762347,22 2,20.9323876 2,19.6153846 L2,16.8461538 C2,15.5291721 3.06762994,14.4615385 4.38461538,14.4615385 L4.8215823,14.4609378 C4.95831893,13.6120029 5.3588057,12.8211623 5.97459937,12.2053686 C6.69125996,11.488708 7.64500941,11.0636656 8.6514968,11.0066017 L8.88461538,11 C9.44565477,11 9.98370225,10.7771334 10.3804178,10.3804178 C10.6184472,10.1423885 10.7938909,9.85347965 10.8961612,9.53925278 L10.6153846,9.53846154 C9.29840448,9.53846154 8.23076923,8.47082629 8.23076923,7.15384615 L8.23076923,4.38461538 C8.23076923,3.06762994 9.29840286,2 10.6153846,2 L13.3846154,2 Z M7.15384615,16.4615385 L4.38461538,16.4615385 C4.17220099,16.4615385 4,16.63374 4,16.8461538 L4,19.6153846 C4,19.8278134 4.17218833,20 4.38461538,20 L7.15384615,20 C7.36626945,20 7.53846154,19.8278103 7.53846154,19.6153846 L7.53846154,16.8461538 C7.53846154,16.6337432 7.36625679,16.4615385 7.15384615,16.4615385 Z M19.6153846,16.4615385 L16.8461538,16.4615385 C16.6337432,16.4615385 16.4615385,16.6337432 16.4615385,16.8461538 L16.4615385,19.6153846 C16.4615385,19.8278103 16.6337306,20 16.8461538,20 L19.6153846,20 C19.8278229,20 20,19.8278229 20,19.6153846 L20,16.8461538 C20,16.6337306 19.8278103,16.4615385 19.6153846,16.4615385 Z M13.3846154,4 L10.6153846,4 C10.4029708,4 10.2307692,4.17220099 10.2307692,4.38461538 L10.2307692,7.15384615 C10.2307692,7.36625679 10.402974,7.53846154 10.6153846,7.53846154 L13.3846154,7.53846154 C13.597026,7.53846154 13.7692308,7.36625679 13.7692308,7.15384615 L13.7692308,4.38461538 C13.7692308,4.17220099 13.5970292,4 13.3846154,4 Z" id="Shape" fill-rule="nonzero"/></g></symbol><symbol id="icon-eds-small-arrow-left" viewBox="0 0 16 17"><path stroke="currentColor" stroke-linecap="round" stroke-linejoin="round" stroke-width="2" d="M14 8.092H2m0 0L8 2M2 8.092l6 6.035"/></symbol><symbol id="icon-eds-small-arrow-right" viewBox="0 0 16 16"><g fill-rule="evenodd" stroke="currentColor" stroke-linecap="round" stroke-linejoin="round" stroke-width="2"><path d="M2 8.092h12M8 2l6 6.092M8 14.127l6-6.035"/></g></symbol><symbol id="icon-orcid-logo" viewBox="0 0 40 40"><path fill-rule="evenodd" d="M12.281 10.453c.875 0 1.578-.719 1.578-1.578 0-.86-.703-1.578-1.578-1.578-.875 0-1.578.703-1.578 1.578 0 .86.703 1.578 1.578 1.578Zm-1.203 18.641h2.406V12.359h-2.406v16.735Z"/><path fill-rule="evenodd" d="M17.016 12.36h6.5c6.187 0 8.906 4.421 8.906 8.374 0 4.297-3.36 8.375-8.875 8.375h-6.531V12.36Zm6.234 14.578h-3.828V14.53h3.703c4.688 0 6.828 2.844 6.828 6.203 0 2.063-1.25 6.203-6.703 6.203Z" clip-rule="evenodd"/></symbol></svg> </div> <a class="c-skip-link" href="#main">Skip to main content</a> <header class="eds-c-header" data-eds-c-header> <div class="eds-c-header__container" data-eds-c-header-expander-anchor> <div class="eds-c-header__brand"> <a href="https://link.springer.com" data-test=springerlink-logo data-track="click_imprint_logo" data-track-context="unified header" data-track-action="click logo link" data-track-category="unified header" data-track-label="link" > <img src="/oscar-static/images/darwin/header/img/logo-springer-nature-link-3149409f62.svg" alt="Springer Nature Link"> </a> </div> <a class="c-header__link eds-c-header__link" id="identity-account-widget" href='https://idp.springer.com/auth/personal/springernature?redirect_uri=https://link.springer.com/article/10.1007/s10539-014-9469-4?'><span class="eds-c-header__widget-fragment-title">Log in</span></a> </div> <nav class="eds-c-header__nav" aria-label="header navigation"> <div class="eds-c-header__nav-container"> <div class="eds-c-header__item eds-c-header__item--menu"> <a href="#eds-c-header-nav" class="eds-c-header__link" data-eds-c-header-expander> <svg class="eds-c-header__icon" width="24" height="24" aria-hidden="true" focusable="false"> <use xlink:href="#icon-eds-i-menu-medium"></use> </svg><span>Menu</span> </a> </div> <div class="eds-c-header__item eds-c-header__item--inline-links"> <a class="eds-c-header__link" href="https://link.springer.com/journals/" data-track="nav_find_a_journal" data-track-context="unified header" data-track-action="click find a journal" data-track-category="unified header" data-track-label="link" > Find a journal </a> <a class="eds-c-header__link" href="https://www.springernature.com/gp/authors" data-track="nav_how_to_publish" data-track-context="unified header" data-track-action="click publish with us link" data-track-category="unified header" data-track-label="link" > Publish with us </a> <a class="eds-c-header__link" href="https://link.springernature.com/home/" data-track="nav_track_your_research" data-track-context="unified header" data-track-action="click track your research" data-track-category="unified header" data-track-label="link" > Track your research </a> </div> <div class="eds-c-header__link-container"> <div class="eds-c-header__item eds-c-header__item--divider"> <a href="#eds-c-header-popup-search" class="eds-c-header__link" data-eds-c-header-expander data-eds-c-header-test-search-btn> <svg class="eds-c-header__icon" width="24" height="24" aria-hidden="true" focusable="false"> <use xlink:href="#icon-eds-i-search-medium"></use> </svg><span>Search</span> </a> </div> <div id="ecommerce-header-cart-icon-link" class="eds-c-header__item ecommerce-cart" style="display:inline-block"> <a class="eds-c-header__link" href="https://order.springer.com/public/cart" style="appearance:none;border:none;background:none;color:inherit;position:relative"> <svg id="eds-i-cart" class="eds-c-header__icon" xmlns="http://www.w3.org/2000/svg" height="24" width="24" viewBox="0 0 24 24" aria-hidden="true" focusable="false"> <path fill="currentColor" fill-rule="nonzero" d="M2 1a1 1 0 0 0 0 2l1.659.001 2.257 12.808a2.599 2.599 0 0 0 2.435 2.185l.167.004 9.976-.001a2.613 2.613 0 0 0 2.61-1.748l.03-.106 1.755-7.82.032-.107a2.546 2.546 0 0 0-.311-1.986l-.108-.157a2.604 2.604 0 0 0-2.197-1.076L6.042 5l-.56-3.17a1 1 0 0 0-.864-.82l-.12-.007L2.001 1ZM20.35 6.996a.63.63 0 0 1 .54.26.55.55 0 0 1 .082.505l-.028.1L19.2 15.63l-.022.05c-.094.177-.282.299-.526.317l-10.145.002a.61.61 0 0 1-.618-.515L6.394 6.999l13.955-.003ZM18 19a2 2 0 1 0 0 4 2 2 0 0 0 0-4ZM8 19a2 2 0 1 0 0 4 2 2 0 0 0 0-4Z"></path> </svg><span>Cart</span><span class="cart-info" style="display:none;position:absolute;top:10px;right:45px;background-color:#C65301;color:#fff;width:18px;height:18px;font-size:11px;border-radius:50%;line-height:17.5px;text-align:center"></span></a> <script>(function () { var exports = {}; if (window.fetch) { "use strict"; Object.defineProperty(exports, "__esModule", { value: true }); exports.headerWidgetClientInit = void 0; var headerWidgetClientInit = function (getCartInfo) { document.body.addEventListener("updatedCart", function () { updateCartIcon(); }, false); return updateCartIcon(); function updateCartIcon() { return getCartInfo() .then(function (res) { return res.json(); }) .then(refreshCartState) .catch(function (_) { }); } function refreshCartState(json) { var indicator = document.querySelector("#ecommerce-header-cart-icon-link .cart-info"); /* istanbul ignore else */ if (indicator && json.itemCount) { indicator.style.display = 'block'; indicator.textContent = json.itemCount > 9 ? '9+' : json.itemCount.toString(); var moreThanOneItem = json.itemCount > 1; indicator.setAttribute('title', "there ".concat(moreThanOneItem ? "are" : "is", " ").concat(json.itemCount, " item").concat(moreThanOneItem ? "s" : "", " in your cart")); } return json; } }; exports.headerWidgetClientInit = headerWidgetClientInit; headerWidgetClientInit( function () { return window.fetch("https://cart.springer.com/cart-info", { credentials: "include", headers: { Accept: "application/json" } }) } ) }})()</script> </div> </div> </div> </nav> </header> <article lang="en" id="main" class="app-masthead__colour-21"> <section class="app-masthead " aria-label="article masthead"> <div class="app-masthead__container"> <div class="app-article-masthead u-sans-serif js-context-bar-sticky-point-masthead" data-track-component="article" data-test="masthead-component"> <div class="app-article-masthead__info"> <nav aria-label="breadcrumbs" data-test="breadcrumbs"> <ol class="c-breadcrumbs c-breadcrumbs--contrast" itemscope itemtype="https://schema.org/BreadcrumbList"> <li class="c-breadcrumbs__item" id="breadcrumb0" itemprop="itemListElement" itemscope="" itemtype="https://schema.org/ListItem"> <a href="/" class="c-breadcrumbs__link" itemprop="item" data-track="click_breadcrumb" data-track-context="article page" data-track-category="article" data-track-action="breadcrumbs" data-track-label="breadcrumb1"><span itemprop="name">Home</span></a><meta itemprop="position" content="1"> <svg class="c-breadcrumbs__chevron" role="img" aria-hidden="true" focusable="false" width="10" height="10" viewBox="0 0 10 10"> <path d="m5.96738168 4.70639573 2.39518594-2.41447274c.37913917-.38219212.98637524-.38972225 1.35419292-.01894278.37750606.38054586.37784436.99719163-.00013556 1.37821513l-4.03074001 4.06319683c-.37758093.38062133-.98937525.38100976-1.367372-.00003075l-4.03091981-4.06337806c-.37759778-.38063832-.38381821-.99150444-.01600053-1.3622839.37750607-.38054587.98772445-.38240057 1.37006824.00302197l2.39538588 2.4146743.96295325.98624457z" fill-rule="evenodd" transform="matrix(0 -1 1 0 0 10)"/> </svg> </li> <li class="c-breadcrumbs__item" id="breadcrumb1" itemprop="itemListElement" itemscope="" itemtype="https://schema.org/ListItem"> <a href="/journal/10539" class="c-breadcrumbs__link" itemprop="item" data-track="click_breadcrumb" data-track-context="article page" data-track-category="article" data-track-action="breadcrumbs" data-track-label="breadcrumb2"><span itemprop="name">Biology & Philosophy</span></a><meta itemprop="position" content="2"> <svg class="c-breadcrumbs__chevron" role="img" aria-hidden="true" focusable="false" width="10" height="10" viewBox="0 0 10 10"> <path d="m5.96738168 4.70639573 2.39518594-2.41447274c.37913917-.38219212.98637524-.38972225 1.35419292-.01894278.37750606.38054586.37784436.99719163-.00013556 1.37821513l-4.03074001 4.06319683c-.37758093.38062133-.98937525.38100976-1.367372-.00003075l-4.03091981-4.06337806c-.37759778-.38063832-.38381821-.99150444-.01600053-1.3622839.37750607-.38054587.98772445-.38240057 1.37006824.00302197l2.39538588 2.4146743.96295325.98624457z" fill-rule="evenodd" transform="matrix(0 -1 1 0 0 10)"/> </svg> </li> <li class="c-breadcrumbs__item" id="breadcrumb2" itemprop="itemListElement" itemscope="" itemtype="https://schema.org/ListItem"> <span itemprop="name">Article</span><meta itemprop="position" content="3"> </li> </ol> </nav> <h1 class="c-article-title" data-test="article-title" data-article-title="">Fish do not feel pain and its implications for understanding phenomenal consciousness</h1> <ul class="c-article-identifiers"> <li class="c-article-identifiers__item"> <a href="https://www.springernature.com/gp/open-research/about/the-fundamentals-of-open-access-and-open-research" data-track="click" data-track-action="open access" data-track-label="link" class="u-color-open-access" data-test="open-access">Open access</a> </li> <li class="c-article-identifiers__item"> Published: <time datetime="2014-12-16">16 December 2014</time> </li> </ul> <ul class="c-article-identifiers c-article-identifiers--cite-list"> <li class="c-article-identifiers__item"> <span data-test="journal-volume">Volume 30</span>, pages 149–165, (<span data-test="article-publication-year">2015</span>) </li> <li class="c-article-identifiers__item c-article-identifiers__item--cite"> <a href="#citeas" data-track="click" data-track-action="cite this article" data-track-category="article body" data-track-label="link">Cite this article</a> </li> </ul> <div class="app-article-masthead__buttons" data-test="download-article-link-wrapper" data-track-context="masthead"> <div class="c-pdf-container"> <div class="c-pdf-download u-clear-both u-mb-16"> <a href="/content/pdf/10.1007/s10539-014-9469-4.pdf" class="u-button u-button--full-width u-button--primary u-justify-content-space-between c-pdf-download__link" data-article-pdf="true" data-readcube-pdf-url="true" data-test="pdf-link" data-draft-ignore="true" data-track="content_download" data-track-type="article pdf download" data-track-action="download pdf" data-track-label="button" data-track-external download> <span class="c-pdf-download__text">Download PDF</span> <svg aria-hidden="true" focusable="false" width="16" height="16" class="u-icon"><use xlink:href="#icon-eds-i-download-medium"/></svg> </a> </div> </div> <p class="app-article-masthead__access"> <svg width="16" height="16" focusable="false" role="img" aria-hidden="true"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-check-filled-medium"></use></svg> You have full access to this <a href="https://www.springernature.com/gp/open-research/about/the-fundamentals-of-open-access-and-open-research" data-track="click" data-track-action="open access" data-track-label="link">open access</a> article</p> </div> </div> <div class="app-article-masthead__brand"> <a href="/journal/10539" class="app-article-masthead__journal-link" data-track="click_journal_home" data-track-action="journal homepage" data-track-context="article page" data-track-label="link"> <picture> <source type="image/webp" media="(min-width: 768px)" width="120" height="159" srcset="https://media.springernature.com/w120/springer-static/cover-hires/journal/10539?as=webp, https://media.springernature.com/w316/springer-static/cover-hires/journal/10539?as=webp 2x"> <img width="72" height="95" src="https://media.springernature.com/w72/springer-static/cover-hires/journal/10539?as=webp" srcset="https://media.springernature.com/w144/springer-static/cover-hires/journal/10539?as=webp 2x" alt=""> </picture> <span class="app-article-masthead__journal-title">Biology & Philosophy</span> </a> <a href="https://link.springer.com/journal/10539/aims-and-scope" class="app-article-masthead__submission-link" data-track="click_aims_and_scope" data-track-action="aims and scope" data-track-context="article page" data-track-label="link"> Aims and scope <svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-arrow-right-medium"></use></svg> </a> <a href="https://www.editorialmanager.com/biph" class="app-article-masthead__submission-link" data-track="click_submit_manuscript" data-track-context="article masthead on springerlink article page" data-track-action="submit manuscript" data-track-label="link"> Submit manuscript <svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-arrow-right-medium"></use></svg> </a> </div> </div> </div> </section> <div class="c-article-main u-container u-mt-24 u-mb-32 l-with-sidebar" id="main-content" data-component="article-container"> <main class="u-serif js-main-column" data-track-component="article body"> <div class="c-context-bar u-hide" data-test="context-bar" data-context-bar aria-hidden="true"> <div class="c-context-bar__container u-container"> <div class="c-context-bar__title"> Fish do not feel pain and its implications for understanding phenomenal consciousness </div> <div data-test="inCoD" data-track-context="sticky banner"> <div class="c-pdf-container"> <div class="c-pdf-download u-clear-both u-mb-16"> <a href="/content/pdf/10.1007/s10539-014-9469-4.pdf" class="u-button u-button--full-width u-button--primary u-justify-content-space-between c-pdf-download__link" data-article-pdf="true" data-readcube-pdf-url="true" data-test="pdf-link" data-draft-ignore="true" data-track="content_download" data-track-type="article pdf download" data-track-action="download pdf" data-track-label="button" data-track-external download> <span class="c-pdf-download__text">Download PDF</span> <svg aria-hidden="true" focusable="false" width="16" height="16" class="u-icon"><use xlink:href="#icon-eds-i-download-medium"/></svg> </a> </div> </div> </div> </div> </div> <div class="c-article-header"> <header> <ul class="c-article-author-list c-article-author-list--short" data-test="authors-list" data-component-authors-activator="authors-list"><li class="c-article-author-list__item"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Brian-Key-Aff1" data-author-popup="auth-Brian-Key-Aff1" data-author-search="Key, Brian" data-corresp-id="c1">Brian Key<svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-mail-medium"></use></svg></a><sup class="u-js-hide"><a href="#Aff1">1</a></sup> </li></ul> <div data-test="article-metrics"> <ul class="app-article-metrics-bar u-list-reset"> <li class="app-article-metrics-bar__item"> <p class="app-article-metrics-bar__count"><svg class="u-icon app-article-metrics-bar__icon" width="24" height="24" aria-hidden="true" focusable="false"> <use xlink:href="#icon-eds-i-accesses-medium"></use> </svg>19k <span class="app-article-metrics-bar__label">Accesses</span></p> </li> <li class="app-article-metrics-bar__item"> <p class="app-article-metrics-bar__count"><svg class="u-icon app-article-metrics-bar__icon" width="24" height="24" aria-hidden="true" focusable="false"> <use xlink:href="#icon-eds-i-citations-medium"></use> </svg>52 <span class="app-article-metrics-bar__label">Citations</span></p> </li> <li class="app-article-metrics-bar__item"> <p class="app-article-metrics-bar__count"><svg class="u-icon app-article-metrics-bar__icon" width="24" height="24" aria-hidden="true" focusable="false"> <use xlink:href="#icon-eds-i-altmetric-medium"></use> </svg>223 <span class="app-article-metrics-bar__label">Altmetric</span></p> </li> <li class="app-article-metrics-bar__item"> <p class="app-article-metrics-bar__count"><svg class="u-icon app-article-metrics-bar__icon app-article-metrics-bar__icon--mentions" width="24" height="24" aria-hidden="true" focusable="false"> <use xlink:href="#icon-eds-i-mentions-medium"></use> </svg>31 <span class="app-article-metrics-bar__label">Mentions</span></p> </li> <li class="app-article-metrics-bar__item app-article-metrics-bar__item--metrics"> <p class="app-article-metrics-bar__details"><a href="/article/10.1007/s10539-014-9469-4/metrics" data-track="click" data-track-action="view metrics" data-track-label="link" rel="nofollow">Explore all metrics <svg class="u-icon app-article-metrics-bar__arrow-icon" width="24" height="24" aria-hidden="true" focusable="false"> <use xlink:href="#icon-eds-i-arrow-right-medium"></use> </svg></a></p> </li> </ul> </div> <div class="u-mt-32"> </div> </header> </div> <div data-article-body="true" data-track-component="article body" class="c-article-body"> <section aria-labelledby="Abs1" data-title="Abstract" lang="en"><div class="c-article-section" id="Abs1-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Abs1">Abstract</h2><div class="c-article-section__content" id="Abs1-content"><p>Phenomenal consciousness or the subjective experience of feeling sensory stimuli is fundamental to human existence. Because of the ubiquity of their subjective experiences, humans seem to readily accept the anthropomorphic extension of these mental states to other animals. Humans will typically extrapolate feelings of pain to animals if they respond physiologically and behaviourally to noxious stimuli. The alternative view that fish instead respond to noxious stimuli reflexly and with a limited behavioural repertoire is defended within the context of our current understanding of the neuroanatomy and neurophysiology of mental states. Consequently, a set of fundamental properties of neural tissue necessary for feeling pain or experiencing affective states in vertebrates is proposed. While mammals and birds possess the prerequisite neural architecture for phenomenal consciousness, it is concluded that fish lack these essential characteristics and hence do not feel pain.</p></div></div></section> <div data-test="cobranding-download"> </div> <section aria-labelledby="inline-recommendations" data-title="Inline Recommendations" class="c-article-recommendations" data-track-component="inline-recommendations"> <h3 class="c-article-recommendations-title" id="inline-recommendations">Similar content being viewed by others</h3> <div class="c-article-recommendations-list"> <div class="c-article-recommendations-list__item"> <article class="c-article-recommendations-card" itemscope itemtype="http://schema.org/ScholarlyArticle"> <div class="c-article-recommendations-card__img"><img src="https://media.springernature.com/w92h120/springer-static/cover-hires/book/978-3-030-31011-0?as=webp" loading="lazy" alt=""></div> <div class="c-article-recommendations-card__main"> <h3 class="c-article-recommendations-card__heading" itemprop="name headline"> <a class="c-article-recommendations-card__link" itemprop="url" href="https://link.springer.com/10.1007/978-3-030-31011-0_6?fromPaywallRec=false" data-track="select_recommendations_1" data-track-context="inline recommendations" data-track-action="click recommendations inline - 1" data-track-label="10.1007/978-3-030-31011-0_6">Speciesism and Human Supremacy in Animal Neuroscience </a> </h3> <div class="c-article-meta-recommendations" data-test="recommendation-info"> <span class="c-article-meta-recommendations__item-type">Chapter</span> <span class="c-article-meta-recommendations__date">© 2020</span> </div> </div> </article> </div> <div class="c-article-recommendations-list__item"> <article class="c-article-recommendations-card" itemscope itemtype="http://schema.org/ScholarlyArticle"> <div class="c-article-recommendations-card__img"><img src="https://media.springernature.com/w92h120/springer-static/cover-hires/book/978-3-030-41675-1?as=webp" loading="lazy" alt=""></div> <div class="c-article-recommendations-card__main"> <h3 class="c-article-recommendations-card__heading" itemprop="name headline"> <a class="c-article-recommendations-card__link" itemprop="url" href="https://link.springer.com/10.1007/978-3-030-41675-1_7?fromPaywallRec=false" data-track="select_recommendations_2" data-track-context="inline recommendations" data-track-action="click recommendations inline - 2" data-track-label="10.1007/978-3-030-41675-1_7">Inside the Fish Brain: Cognition, Learning and Consciousness </a> </h3> <div class="c-article-meta-recommendations" data-test="recommendation-info"> <span class="c-article-meta-recommendations__item-type">Chapter</span> <span class="c-article-meta-recommendations__date">© 2020</span> </div> </div> </article> </div> <div class="c-article-recommendations-list__item"> <article class="c-article-recommendations-card" itemscope itemtype="http://schema.org/ScholarlyArticle"> <div class="c-article-recommendations-card__img"><img src="https://media.springernature.com/w92h120/springer-static/cover-hires/book/978-3-662-44866-3?as=webp" loading="lazy" alt=""></div> <div class="c-article-recommendations-card__main"> <h3 class="c-article-recommendations-card__heading" itemprop="name headline"> <a class="c-article-recommendations-card__link" itemprop="url" href="https://link.springer.com/10.1007/7854_2014_278?fromPaywallRec=false" data-track="select_recommendations_3" data-track-context="inline recommendations" data-track-action="click recommendations inline - 3" data-track-label="10.1007/7854_2014_278">A Framework for Investigating Animal Consciousness </a> </h3> <div class="c-article-meta-recommendations" data-test="recommendation-info"> <span class="c-article-meta-recommendations__item-type">Chapter</span> <span class="c-article-meta-recommendations__date">© 2015</span> </div> </div> </article> </div> </div> </section> <script> window.dataLayer = window.dataLayer || []; window.dataLayer.push({ recommendations: { recommender: 'semantic', model: 'specter', policy_id: 'NA', timestamp: 1732838534, embedded_user: 'null' } }); </script> <div class="app-card-service" data-test="article-checklist-banner"> <div> <a class="app-card-service__link" data-track="click_presubmission_checklist" data-track-context="article page top of reading companion" data-track-category="pre-submission-checklist" data-track-action="clicked article page checklist banner test 2 old version" data-track-label="link" href="https://beta.springernature.com/pre-submission?journalId=10539" data-test="article-checklist-banner-link"> <span class="app-card-service__link-text">Use our pre-submission checklist</span> <svg class="app-card-service__link-icon" aria-hidden="true" focusable="false"><use xlink:href="#icon-eds-i-arrow-right-small"></use></svg> </a> <p class="app-card-service__description">Avoid common mistakes on your manuscript.</p> </div> <div class="app-card-service__icon-container"> <svg class="app-card-service__icon" aria-hidden="true" focusable="false"> <use xlink:href="#icon-eds-i-clipboard-check-medium"></use> </svg> </div> </div> <div class="main-content"> <section data-title="Introduction"><div class="c-article-section" id="Sec1-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec1">Introduction</h2><div class="c-article-section__content" id="Sec1-content"><p>There is a belief in some scientific and lay communities that because fish respond behaviourally to noxious stimuli, then <i>ipso facto</i>, fish feel pain. Sneddon (<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2011" title="Sneddon LU (2011) Nociception or pain in fish. In: Farrell A (ed) Encyclopedia of fish physiology. Academic Press, London, pp 713–719. ISBN 978-0-12-374545-3" href="/article/10.1007/s10539-014-9469-4#ref-CR113" id="ref-link-section-d30854801e292">2011</a>) clearly articulates the logic by stating: “to explore the possibility of pain perception in nonhumans we use indirect measures similar to those used for human infants who cannot convey whether they are in pain. We measure physiological responses (e.g., cardiovascular) and behavioral changes (e.g. withdrawal) to assess whether a tissue-damaging event is painful to an animal”. In some cases, the inference that fish have affective states arises because of conflation of nociception with pain (Demski <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2013" title="Demski LS (2013) The pallium and mind/behavior relationships in teleost fishes. Brain Behav Evol 82:31–44" href="/article/10.1007/s10539-014-9469-4#ref-CR23" id="ref-link-section-d30854801e295">2013</a>; Kittilsen <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2013" title="Kittilsen S (2013) Functional aspects of emotions in fish. Behav Process 100:153–159" href="/article/10.1007/s10539-014-9469-4#ref-CR130" id="ref-link-section-d30854801e298">2013</a>; Malafoglia et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2013" title="Malafoglia V, Bryant B, Raffaeli W, Giordano A, Bellipanni G (2013) The zebrafish as a model for nociception studies. J Cell Physiol 228:1956–1966" href="/article/10.1007/s10539-014-9469-4#ref-CR62" id="ref-link-section-d30854801e301">2013</a>). Interestingly, sometimes the difference between nociception and pain is recognized but it is still considered safer to err on the side of caution and accept that fish feel pain (Jones <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2013" title="Jones RC (2013) Science, sentience, and animal welfare. Biol Philos 28:1–30" href="/article/10.1007/s10539-014-9469-4#ref-CR45" id="ref-link-section-d30854801e305">2013</a>). Unfortunately, endowing fish with the subjective ability to experience pain is typically undertaken without reference to its neurophysiological bases (Rose <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2002" title="Rose JD (2002) The neurobiological nature of fishes and the question of awareness and pain. Rev Fish Sci 10:1–38" href="/article/10.1007/s10539-014-9469-4#ref-CR98" id="ref-link-section-d30854801e308">2002</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2007" title="Rose JD (2007) Anthropomorphism and ‘mental welfare’ of fishes. Dis Aquat Organ 75:139–154" href="/article/10.1007/s10539-014-9469-4#ref-CR99" id="ref-link-section-d30854801e311">2007</a>; Browman and Skiftesvik <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2011" title="Browman HI, Skiftesvik AB (2011) Welfare in aquatic organisms—is there some faith-based HAR-King going on here? Dis Aquat Org 94:255–257" href="/article/10.1007/s10539-014-9469-4#ref-CR13" id="ref-link-section-d30854801e314">2011</a>).</p><p>Before interrogating the issue of fish feeling pain and its implications for phenomenal consciousness, I will briefly define several key terms. When I refer to fish it is with the knowledge that this is a highly diverse paraphyletic group consisting of ~30,000 species. Since most of the behavioural and neuroanatomical investigations discussed here have been undertaken only on a small number of ray-finned fish, there is considerable extrapolation involved when I use the generic term fish. A noxious stimulus is one that is considered to be physically harmful to an animal without reference to feelings. For example, excessive heat, a skin incision, toxic chemical exposure and extreme mechanical pressure are all stimuli that can perturb normal tissue morphology, and are hence considered to be noxious. Nociception is referred to as the neurobiological processes associated with the activation of peripheral sensory neurons and their upstream neural pathways by noxious stimuli in the absence of conscious feeling. In contrast, pain is the subjective experience of feeling a noxious stimulus (however, in certain central neuropathies in humans it can arise without external stimuli). The subjective “feeling” associated with a sensory stimulus is also referred to as a “quale” or “phenomenal consciousness” (Kanai and Tsuchiya <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2012" title="Kanai R, Tsuchiya N (2012) Qualia. Curr Biol 22:R392–R396" href="/article/10.1007/s10539-014-9469-4#ref-CR50" id="ref-link-section-d30854801e320">2012</a>). Given the above, I acknowledge the tautology in the manuscript’s title since the word “pain” is already defined as “to feel a noxious stimuli”. However, the phrase “feel pain” within the title was chosen to over-emphasize the subjective or qualitative nature of pain.</p><p>One of the main proponents in the literature of the thesis that fish do not feel pain has been John D. Rose. In a series of comprehensive articles (Rose <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2002" title="Rose JD (2002) The neurobiological nature of fishes and the question of awareness and pain. Rev Fish Sci 10:1–38" href="/article/10.1007/s10539-014-9469-4#ref-CR98" id="ref-link-section-d30854801e326">2002</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2007" title="Rose JD (2007) Anthropomorphism and ‘mental welfare’ of fishes. Dis Aquat Organ 75:139–154" href="/article/10.1007/s10539-014-9469-4#ref-CR99" id="ref-link-section-d30854801e329">2007</a>; Rose et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2014" title="Rose JD, Arlinghaus R, Cooke SJ, Diggles BK, Sawynok W, Stevens ED, Wynne CDL (2014) Can fish really feel pain? Fish Fish 15:97–133" href="/article/10.1007/s10539-014-9469-4#ref-CR100" id="ref-link-section-d30854801e332">2014</a>) it was argued that fish do not experience the sensation of pain. Anthropomorphism was considered as a hindrance to understanding the underlying causes of behavioural responses of animals to sensory stimuli (Rose <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2002" title="Rose JD (2002) The neurobiological nature of fishes and the question of awareness and pain. Rev Fish Sci 10:1–38" href="/article/10.1007/s10539-014-9469-4#ref-CR98" id="ref-link-section-d30854801e335">2002</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2007" title="Rose JD (2007) Anthropomorphism and ‘mental welfare’ of fishes. Dis Aquat Organ 75:139–154" href="/article/10.1007/s10539-014-9469-4#ref-CR99" id="ref-link-section-d30854801e338">2007</a>). Rose advocated attention to the evolution, development and organization of the nervous system in order to understand fish behaviour. He initially drew attention to three key issues (Rose <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2002" title="Rose JD (2002) The neurobiological nature of fishes and the question of awareness and pain. Rev Fish Sci 10:1–38" href="/article/10.1007/s10539-014-9469-4#ref-CR98" id="ref-link-section-d30854801e342">2002</a>). First, behavioural responses to sensory stimuli must be distinguished from psychological experiences. Second, the cerebral cortex in humans is fundamental for the awareness of sensory stimuli. Third, fish lack a cerebral cortex or its homologue and hence cannot experience pain or fear. In 2007, Rose highlighted the problems of anthropomorphic thinking in respect to fish behaviour and how it influenced welfare issues. He stressed that pain and emotion were not primitive feelings that arose early in vertebrate evolution but were rather more recent acquisitions, associated with the emergence of the cerebral cortex (Rose <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2007" title="Rose JD (2007) Anthropomorphism and ‘mental welfare’ of fishes. Dis Aquat Organ 75:139–154" href="/article/10.1007/s10539-014-9469-4#ref-CR99" id="ref-link-section-d30854801e345">2007</a>). In 2014, Rose et al. (<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2014" title="Rose JD, Arlinghaus R, Cooke SJ, Diggles BK, Sawynok W, Stevens ED, Wynne CDL (2014) Can fish really feel pain? Fish Fish 15:97–133" href="/article/10.1007/s10539-014-9469-4#ref-CR100" id="ref-link-section-d30854801e348">2014</a>) rebutted experimental evidence supposedly supporting claims that fish feel pain. They demonstrated deficiencies in methodological approaches and highlighted problems in concluding pain experience from behavioural responses. Moreover, they recognized that teleosts typically lack nociceptors responsible for transmission of pain but instead have an abundance of A-delta fibres that are most likely subserving escape and avoidance responses rather than the experience of pain.</p><p>Despite the work of Rose et al. (Rose <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2002" title="Rose JD (2002) The neurobiological nature of fishes and the question of awareness and pain. Rev Fish Sci 10:1–38" href="/article/10.1007/s10539-014-9469-4#ref-CR98" id="ref-link-section-d30854801e354">2002</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2007" title="Rose JD (2007) Anthropomorphism and ‘mental welfare’ of fishes. Dis Aquat Organ 75:139–154" href="/article/10.1007/s10539-014-9469-4#ref-CR99" id="ref-link-section-d30854801e357">2007</a>; Rose et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2014" title="Rose JD, Arlinghaus R, Cooke SJ, Diggles BK, Sawynok W, Stevens ED, Wynne CDL (2014) Can fish really feel pain? Fish Fish 15:97–133" href="/article/10.1007/s10539-014-9469-4#ref-CR100" id="ref-link-section-d30854801e360">2014</a>) there remains a strong trend in the literature to bestow fish with the ability to feel pain and to experience fear and other emotions. The alternate view that fish do not feel pain or experience affective states needs more careful consideration, particularly as it has consequences for understanding the neuroanatomical basis of phenomenal consciousness. Here I consolidate the arguments for why fish are believed to feel pain into six main reasons. By undertaking a deeper analysis of the behavioural observations in the light of our understanding of neurophysiology and neuroanatomy, I subsequently propose that it is more plausible and probable to reason that fish do not feel pain. Concluding that fish do not feel pain affords an opportunity to define the basic architectural properties of the neural circuitry necessary for phenomenal consciousness through comparisons of fish and mammalian neuroanatomies. These properties then provide a simple tool for assessing the likelihood that a vertebrate animal will experience “feelings” such as pain.</p></div></div></section><section data-title="What are the reasons for the anthropomorphic view that fish feel pain?"><div class="c-article-section" id="Sec2-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec2">What are the reasons for the anthropomorphic view that fish feel pain?</h2><div class="c-article-section__content" id="Sec2-content"><p>There are six principal reasons that account for why some people believe that fish feel pain. One, fish demonstrate behaviours consistent with the way humans might react to noxious stimuli that cause pain. For example, fish will either attempt to rapidly escape or display anomalous behaviour (Reilly et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2008" title="Reilly SC, Quinn JP, Cossins AR, Sneddon LU (2008) Behavioural analysis of a nociceptive event in fish: comparisons between three species demonstrate specific responses. Appl Anim Behav Sci 114:248–259" href="/article/10.1007/s10539-014-9469-4#ref-CR95" id="ref-link-section-d30854801e371">2008</a>) in response to noxious stimuli, such as electric shock or a chemical irritant. Two, medicating fish with an analgesic (a drug that attenuates pain in humans) reduces the escape response to electric shock (Sneddon <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2003" title="Sneddon LU (2003) The evidence for pain in fish: the use of morphine as an analgesic. Appl Anim Behave Sci 83:153–162" href="/article/10.1007/s10539-014-9469-4#ref-CR111" id="ref-link-section-d30854801e374">2003</a>; Sneddon et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2003" title="Sneddon LU, Braithwaite VA, Gentle MJ (2003) Do fishes have nociceptors? Evidence for the evolution of a vertebrate sensory system. Proc R Soc Lond B 270:1115–1121" href="/article/10.1007/s10539-014-9469-4#ref-CR114" id="ref-link-section-d30854801e377">2003</a>; Jones et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2012" title="Jones SG, Kamunde C, Lemke K, Stevens ED (2012) The dose-response relation for the antinociceptive effect of morphine in a fish, rainbow trout. J Vet Pharmacol Ther 35:563–570" href="/article/10.1007/s10539-014-9469-4#ref-CR47" id="ref-link-section-d30854801e380">2012</a>). Three, fish display classic physiological indicators of stress such as increased ventilation and heart rate and elevated blood levels of the stress hormone cortisol during and after exposure to supposedly stressful stimuli (Reilly et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2008" title="Reilly SC, Quinn JP, Cossins AR, Sneddon LU (2008) Behavioural analysis of a nociceptive event in fish: comparisons between three species demonstrate specific responses. Appl Anim Behav Sci 114:248–259" href="/article/10.1007/s10539-014-9469-4#ref-CR95" id="ref-link-section-d30854801e383">2008</a>; Filk et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2006" title="Filk G, Klaren PHM, Van den Burg EH, Metz JR, Huising MO (2006) CRF and stress in fish. Gen Comp Endocrinol 146:36–44" href="/article/10.1007/s10539-014-9469-4#ref-CR31" id="ref-link-section-d30854801e387">2006</a>; Wolkers et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2013" title="Wolkers CPB, Junior BA, Menescal-de-Oliveira L, Hoffmann A (2013) Stress-induced antinociception in fish reversed by naloxone. PLoS ONE 8:e71175" href="/article/10.1007/s10539-014-9469-4#ref-CR125" id="ref-link-section-d30854801e390">2013</a>). Four, fish have nociceptive nerve fibres and have increased neural activity in the spinal cord, hindbrain and pallium that is specifically associated with a noxious stimulus (Dunlop and Laming <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2005" title="Dunlop R, Laming P (2005) Mechanoreceptive and nociceptive responses in the central nervous system of goldfish (Carassius auratus) and trout (Oncorrhynchus mykiss). J Pain 6:561–568" href="/article/10.1007/s10539-014-9469-4#ref-CR26" id="ref-link-section-d30854801e393">2005</a>). Five, fish can be trained to associate a neutral signal with an impending noxious stimulus and so learn to escape prior to experiencing the noxious stimuli (Dunlop et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2006" title="Dunlop R, Millsopp S, Laming P (2006) Avoidance learning in goldfish (Carassius auratus) and implications for pain perception. Appl Anim Behav Sci 976:255–271" href="/article/10.1007/s10539-014-9469-4#ref-CR27" id="ref-link-section-d30854801e396">2006</a>). Six, it is evolutionarily advantageous to feel pain in order to prevent body injury.</p><h3 class="c-article__sub-heading" id="Sec3">Behavioural responses to noxious stimuli are not necessarily evidence of pain</h3><p>It is common to attribute inner mental states or feelings to organisms or even inanimate objects on the basis of observed behavior. When a noxious stimulus is applied either to the plantar surface of the human foot, or directly to the nerves innervating this region, there is a reflex withdrawal of the lower limb involving contraction of the hip and knee flexors, and relaxation of the extensors. This reflex is protective and enables the rapid removal of the limb from a harmful stimulus. Complete spinal cord injury patients, who lack sensations arising from the lower limb, continue to exhibit the withdrawal flexion reflex (Dimitrijevic and Nathan <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1968" title="Dimitrijevic MR, Nathan PW (1968) Studies of spasticity in man. 3. Analysis of reflex activity evoked by noxious cutaneous stimulation. Brain 91:349–368" href="/article/10.1007/s10539-014-9469-4#ref-CR24" id="ref-link-section-d30854801e406">1968</a>). Thus, reflexes are neither good evidence for, nor a measure of feeling pain. Nonetheless, simple reflex behaviours in response to noxious stimuli continue to be inappropriately used to suggest that fish feel pain.</p><p>Fish exhibit behavioural responses to somatosensory stimulation from a very early stage of development. For example, within the first few days of fertilisation, zebrafish embryos response to touch by initially exhibiting a twitch of their tail, and then slightly later in development, by a few strokes of their tail that elicits a short burst of swimming. While it is tempting to attribute feelings to these embryos, it must be remembered that the telencephalon is not yet morphologically distinct when the touch response first appears at around 21 h post-fertilisation (Hjorth and Key <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2001" title="Hjorth JT, Key B (2001) Are pioneer axons guided by regulatory gene expression domains in the zebrafish forebrain? High-resolution analysis of the patterning of the zebrafish brain during axon tract formation. Dev Biol 229:271–286" href="/article/10.1007/s10539-014-9469-4#ref-CR40" id="ref-link-section-d30854801e412">2001</a>; Saint-Amant <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2006" title="Saint-Amant L (2006) Development of motor networks in zebrafish embryos. Zebrafish 3:173–190" href="/article/10.1007/s10539-014-9469-4#ref-CR102" id="ref-link-section-d30854801e415">2006</a>). Moreover, a lesion to the anterior spinal cord, that isolates the cord from the brain, does not affect the execution of the touch-induced swimming escape response (Pietri et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2009" title="Pietri T, Manalo E, Ryan J, Saint-Amant L, Washbourne P (2009) Glutamate drives the touch response through a rostral loop in the spinal cord of zebrafish embryos. Dev Neurobiol 69:780–795" href="/article/10.1007/s10539-014-9469-4#ref-CR86" id="ref-link-section-d30854801e418">2009</a>). Thus, simple reflex escape behaviours of fish that can be activated by somatosensory stimuli are best not used as evidence for fish experiencing phenomenal consciousness.</p><p>It is important here to draw attention to the fact that pain in humans arises in the forebrain, and is distinct from unconscious behavioural responses mediated by lower brain levels. The forebrain also plays an essential role in pain perception in other mammals. This is elegantly illustrated in a rat model of pain that uses injection of a dilute solution of formalin into the paw. This chemical irritant induces a variety of body movements such as paw shaking, licking and grooming. Animals also exhibit a protective response and attempt to reduce contact of the affected limb with the floor. These behaviours are sometimes considered as indicators of pain. However, rats continued to exhibit such behavioural responses following surgical decerebration (Matthies and Franklin <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1992" title="Matthies BK, Franklin KBJ (1992) Formalin pain is expressed in decerebrate rats but not attenuated by morphine. Pain 51:199–206" href="/article/10.1007/s10539-014-9469-4#ref-CR66" id="ref-link-section-d30854801e424">1992</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1995" title="Matthies BK, Franklin KBJ (1995) Effects of partial decortication on the opioid analgesia in the formalin test. Behav Brain Res 67:59–66" href="/article/10.1007/s10539-014-9469-4#ref-CR67" id="ref-link-section-d30854801e427">1995</a>). One interpretation of these results is that pain is actually experienced in the brainstem, and not in the forebrain in rats. However, this is most unlikely given that systemic administration of an analgesic (morphine) does not attenuate behavioural responses to formalin in decerebrated animals. Morphine was only effective in inhibiting behaviours when connections between the forebrain and brainstem were left intact in sham-operated rats (Matthies and Franklin <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1992" title="Matthies BK, Franklin KBJ (1992) Formalin pain is expressed in decerebrate rats but not attenuated by morphine. Pain 51:199–206" href="/article/10.1007/s10539-014-9469-4#ref-CR66" id="ref-link-section-d30854801e430">1992</a>). Moreover, local application of morphine into either the somatosensory, prefrontal orbital or agranular insular cortices attenuates behavioural responses in the formalin pain model in rats (Soto-Moyano et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1988" title="Soto-Moyano R, Galvez J, Vallejos C, Hernandez A (1988) Topical application of morphine to the rat somatosensory cortex produces analgesia to tonic pain. J Neurosci Res 19:511–514" href="/article/10.1007/s10539-014-9469-4#ref-CR115" id="ref-link-section-d30854801e433">1988</a>; Xie et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2004" title="Xie YF, Wang J, Huo FQ, Jia H, Tang JS (2004) μ but not δ and κ opioid receptor involvement in ventrolateral orbital cortex opioid-evoked antinociception in formalin test rats. Neuroscience 126:717–726" href="/article/10.1007/s10539-014-9469-4#ref-CR126" id="ref-link-section-d30854801e436">2004</a>). Thus, morphine is active in the rat forebrain, which is consistent with it modulating the subjective experience of the noxious stimuli, as in humans (Jones et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1991" title="Jones AKP, Friston KJ, Qi LY, Harris M, Cunningham VJ, Jones T, Feinman C, Frackowiak RSJ (1991) Sites of action of morphine in the brain. Lancet 338:825" href="/article/10.1007/s10539-014-9469-4#ref-CR46" id="ref-link-section-d30854801e440">1991</a>; Taylor et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2013" title="Taylor JJ, Borckardt JJ, Canterberyy M, Li X, Hanlon CA, Brown TR, George MS (2013) Naloxone-reversible modulation of pain circuitry by left prefrontal rTMS. Neuropsychopharmacology 38:1189–1197" href="/article/10.1007/s10539-014-9469-4#ref-CR117" id="ref-link-section-d30854801e443">2013</a>).</p><p>Fish are known to swim away from noxious electric shock and this behavioural response has been used to indicate that these animals feel pain. However, this interpretation is simplistic and can be dismissed given the extensive evidence that fish continue to exhibit escape behaviour following ablation of the entire telencephalon (Hainsworth et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1967" title="Hainsworth FR, Overmier JB, Snowdon CT (1967) Specific and permanent deficits in instrumental avoidance responding following forebrain ablation in the goldfish. J Comp Physiol Psychol 63:111–116" href="/article/10.1007/s10539-014-9469-4#ref-CR39" id="ref-link-section-d30854801e449">1967</a>; Davis et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1976" title="Davis RE, Kassel J, Schwagmeyer P (1976) Telencephalic lesions and behavior in the teleost, macropodus opercularis: reproductive, startle reaction, and operant behavior in the male. Behav Biol 18:165–177" href="/article/10.1007/s10539-014-9469-4#ref-CR22" id="ref-link-section-d30854801e452">1976</a>). Forebrainless fish display no clear evidence of deficits in normal behaviours. For example, forebrainless fish continue to flee from capture by a small fish net with similar locomotor agility as their unoperated counterparts (Kaplan and Aronson <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1967" title="Kaplan H, Aronson LR (1967) Effect of forebrain ablation on the performance of a conditioned avoidance response in the teleost fish, Tilapia H. Macrocephala. Anim Behav 15:438–448" href="/article/10.1007/s10539-014-9469-4#ref-CR51" id="ref-link-section-d30854801e455">1967</a>). The ability to escape or respond to an electric shock is unaffected by removal of either the forebrain or telencephalon in goldfish (Hainsworth et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1967" title="Hainsworth FR, Overmier JB, Snowdon CT (1967) Specific and permanent deficits in instrumental avoidance responding following forebrain ablation in the goldfish. J Comp Physiol Psychol 63:111–116" href="/article/10.1007/s10539-014-9469-4#ref-CR39" id="ref-link-section-d30854801e458">1967</a>; Savage <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1969" title="Savage GE (1969) Some preliminary observations on the role of the telencephalon in food-reinforced behaviour in the goldfish, Carassius auratus. Anim Behav 17:760–772" href="/article/10.1007/s10539-014-9469-4#ref-CR104" id="ref-link-section-d30854801e461">1969</a>; Portavella et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2004a" title="Portavella M, Torres B, Salas C (2004a) Avoidance response in goldfish: emotional and temporal involvement of medial and lateral telencephalic pallium. J Neurosci 24:2335–2342" href="/article/10.1007/s10539-014-9469-4#ref-CR90" id="ref-link-section-d30854801e465">2004a</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference b" title="Portavella M, Torres B, Salas C, Papini MR (2004b) Lesions of the medial pallium, but not of the lateral pallium, disrupt spaced-trial avoidance learning in goldfish (Carassius auratus). Neurosci Lett 362:75–78" href="/article/10.1007/s10539-014-9469-4#ref-CR91" id="ref-link-section-d30854801e468">b</a>) or telencephalon in <i>Tilapia mossambica</i> (Overmier and Gross <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1974" title="Overmier JB, Gross D (1974) Effects of telencephalic ablation upon nest-building and avoidance bahaviors in East African mouthbreeding fish, Tilapia mossambica. Behav Biol 12:211–222" href="/article/10.1007/s10539-014-9469-4#ref-CR82" id="ref-link-section-d30854801e474">1974</a>).</p><p>In summary, the idea that fish flee noxious stimuli because they experience phenomenal consciousness (feel pain) is not the best explanation for this behaviour. It is more probable that fish demonstrate these behaviours because they have evolved innate reflexes associated with specific spinal and sub-telencephalic neural circuits.</p><h3 class="c-article__sub-heading" id="Sec4">Modification of behaviour with drugs does not necessarily demonstrate pain</h3><p>It has been proposed that if an animal’s behavioural response to a noxious stimulus is attenuated following administration of a drug known to be an analgesic in humans, then it is likely that the animal can feel pain. However, it needs to be pointed out that analgesics can be active at multiple sites in the neuroanatomical pathways associated with noxious stimuli. If an analgesic blocks or reduces neural activity in the spinal cord (Yaksh and Rudy <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1976" title="Yaksh TL, Rudy TA (1976) Analgesia mediated by a direct spinal action of narcotics. Science 192:1357–1358" href="/article/10.1007/s10539-014-9469-4#ref-CR128" id="ref-link-section-d30854801e488">1976</a>) it can subsequently attenuate neural responses in the brainstem and telencephalon. Similarly, if an analgesic works at the level of the brainstem it can modulate both brainstem and higher-order brain responses (Pert and Yaksh <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1975" title="Pert A, Yaksh T (1975) Localization of the antinociceptive action of morphine in primate brain. Pharmacol Biochem Behav 3:133–138" href="/article/10.1007/s10539-014-9469-4#ref-CR85" id="ref-link-section-d30854801e491">1975</a>). If an analgesic is active at the level of the telencephalon and reduces behavioural responses (Xie et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2004" title="Xie YF, Wang J, Huo FQ, Jia H, Tang JS (2004) μ but not δ and κ opioid receptor involvement in ventrolateral orbital cortex opioid-evoked antinociception in formalin test rats. Neuroscience 126:717–726" href="/article/10.1007/s10539-014-9469-4#ref-CR126" id="ref-link-section-d30854801e494">2004</a>) then the animal, at least, has the possibility of feeling a noxious stimulus as painful (however this interpretation is dependent first, on the behaviour being non-reflexive and second, on the existence of the necessary neural hardware; see below). At present, the inference that fish feel pain because behavioural responses to noxious stimuli are attenuated following systemic administration of morphine (Sneddon <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2003" title="Sneddon LU (2003) The evidence for pain in fish: the use of morphine as an analgesic. Appl Anim Behave Sci 83:153–162" href="/article/10.1007/s10539-014-9469-4#ref-CR111" id="ref-link-section-d30854801e497">2003</a>) is weak, particularly given that both the site of action as well as the physiological role of this drug in fish are unknown.</p><h3 class="c-article__sub-heading" id="Sec5">Physiological stress is not pain</h3><p>Physiological stress as determined by plasma cortisol levels and opercula beat rate have been used as indicators of feeling pain by fish (Chandroo et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2004" title="Chandroo KP, Duncan IJH, Moccia RD (2004) Can fish suffer? Perspectives on sentience, pain, fear and stress. Appl Anim Behav Sci 86:225–250" href="/article/10.1007/s10539-014-9469-4#ref-CR18" id="ref-link-section-d30854801e508">2004</a>; Braithwaite and Boulcott <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2007" title="Braithwaite VA, Boulcott P (2007) Pain perception, aversion and fear in fish. Dis Aquat Org 75:131–138" href="/article/10.1007/s10539-014-9469-4#ref-CR11" id="ref-link-section-d30854801e511">2007</a>; Scott Weber <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2011" title="Scott Weber E III (2011) Fish analgesia: pain, stress, fear aversion or nociception? Vet Clin Exot Anim 14:21–32" href="/article/10.1007/s10539-014-9469-4#ref-CR106" id="ref-link-section-d30854801e514">2011</a>). The underlying assumption in these cases is that if a fish is exposed to a stimulus that triggers both increased cortisol and behavioural responses, then that fish must be consciously feeling that stimulus as a mental state such as fear and/or pain. If pain was felt by a fish exposed to a physiological stressor, and cortisol was an indicator of the level of discomfort that fish experienced, then one would predict increased cortisol levels in fish as a noxious stimulus was increased. However, this does not appear to be the case. There is no relationship between the apparent “stressful” stimulus and the level of cortisol in fish (Roques et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2010" title="Roques JAC, Abbink W, Geurds F, van de Vis H, Flik G (2010) Tailfin clipping, a painful procedure: studies on Nile tilapia and common carp. Physiol Behav 101:533–540" href="/article/10.1007/s10539-014-9469-4#ref-CR97" id="ref-link-section-d30854801e517">2010</a>). Even when the stimulus causes increased behavioural responses there was no relationship to the level of plasma cortisol. The cortisol response to increased stress seems to be highly variable (Fatira et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2014" title="Fatira E, Papandroulakis N, Pavlidis M (2014) Diel changes in plasma cortisol and effects of size and stress duration on the cortisol response in European sea bass (Dicentrarchus labrax). Fish Physiol Biochem. doi:
 10.1007/s10695-013-9896-1
 
 
 " href="/article/10.1007/s10539-014-9469-4#ref-CR30" id="ref-link-section-d30854801e520">2014</a>; Quillet et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2014" title="Quillet E, Krieg F, Dechamp N, Hervet C, Bérard A, Le Roy P, Guyomard R, Prunet P, Pottinger TG (2014) Quantitative trait loci for magnitude of the plasma cortisol response to confinement in rainbow trout. Genet, Anim. doi:
 10.1111/age.12126
 
 
 " href="/article/10.1007/s10539-014-9469-4#ref-CR93" id="ref-link-section-d30854801e524">2014</a>) and context specific (Manek et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2014" title="Manek AK, Ferrari MC, Niyogi S, Chivers DP (2014) The interactive effects of multiple stressors on physiological stress responders and club cell investment in fathead minnows. Sci Total Environ 467–477:90–97" href="/article/10.1007/s10539-014-9469-4#ref-CR64" id="ref-link-section-d30854801e527">2014</a>). Surprisingly, exposure to multiple stressors simultaneously can lead to decreased rather than an expected increase in cortisol levels (Manek et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2014" title="Manek AK, Ferrari MC, Niyogi S, Chivers DP (2014) The interactive effects of multiple stressors on physiological stress responders and club cell investment in fathead minnows. Sci Total Environ 467–477:90–97" href="/article/10.1007/s10539-014-9469-4#ref-CR64" id="ref-link-section-d30854801e530">2014</a>). Thus, changes in cortisol levels in fish are better explained by autonomic responses to external environmental stresses rather than by internally generated mental states such as fear or pain.</p><h3 class="c-article__sub-heading" id="Sec6">Brain activity in response to noxious activity is not equivalent to pain</h3><p>It has been proposed that fish can feel pain both because they have peripheral nociceptors and because neural responses to noxious stimuli have been recorded in the spinal cord, cerebellum, tectum and telencephalon of fish (Sneddon <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2004" title="Sneddon LU (2004) Evolution of nociception in vertebrates: comparative analysis of lower vertebrates. Brain Res Rev 46:123–130" href="/article/10.1007/s10539-014-9469-4#ref-CR112" id="ref-link-section-d30854801e542">2004</a>; Dunlop and Laming <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2005" title="Dunlop R, Laming P (2005) Mechanoreceptive and nociceptive responses in the central nervous system of goldfish (Carassius auratus) and trout (Oncorrhynchus mykiss). J Pain 6:561–568" href="/article/10.1007/s10539-014-9469-4#ref-CR26" id="ref-link-section-d30854801e545">2005</a>). Nordgreen et al. (<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2007" title="Nordgreen J, Horsberg TE, Ranheim B, Chen ACN (2007) Somatosensory evoked potentials in the telencephalon of Atlanic salmon (Salmo salar) following galvanic stimulation of the tail. J Comp Physiol A 193:1235–1242" href="/article/10.1007/s10539-014-9469-4#ref-CR75" id="ref-link-section-d30854801e548">2007</a>) reported neural activity in the telencephalon following electrical stimulation of the tail of Atlanic salmon. While these authors indicated that this activity is a necessary prerequisite for feeling pain, they realised that it does not necessarily provide evidence for the ability of fish to feel pain. Unfortunately, the neuroanatomical localisation of electrical activity recorded in the telencephalon has not been described. If activity was recorded in the dorsal pallium (homologous to the neocortex; Mueller and Wullimann <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2009" title="Mueller T, Wullimann MF (2009) An evolutionary interpretation of teleostean forebrain anatomy. Brain Behav Evol 74:30–42" href="/article/10.1007/s10539-014-9469-4#ref-CR71" id="ref-link-section-d30854801e551">2009</a>; Mueller et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2011" title="Mueller T, Dong Z, Berberoglu MA, Guo S (2011) The dorsal pallium in zebrafish, Danio rerio (Cyprinidae, Teleostei). Brain Res 1381:95–105" href="/article/10.1007/s10539-014-9469-4#ref-CR72" id="ref-link-section-d30854801e554">2011</a>) of the telencephalon, it would, at least, provide some phylogenetic insight into neural pathways underlying nociception. It would not, however, be evidence of pain or emotion.</p><h3 class="c-article__sub-heading" id="Sec7">Associative learning using noxious stimuli is possible without feeling pain</h3><p>Considering the problems with using simple behavioural responses to noxious stimuli as a measure of pain sensation, avoidance learning has instead been adopted as a means for assessing pain in animals. Rats can easily learn to avoid locations in a cage where electric shocks are delivered and to push a lever that terminates the shock. This learning is viewed as requiring the animal to initially decipher the stimulus (i.e. feeling the stimulus as painful and assessing the intensity using the cerebral cortex; Baastrup et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2010" title="Baastrup C, Maersk-Moller CC, Nyengaard JR, Jensen TS, Finnerup NB (2010) Spinal-, brainstem- and cerebrally mediated responses at- and below-level of a spinal cord contusion in rats: evaluation of pain-like behavior. Pain 151:670–679" href="/article/10.1007/s10539-014-9469-4#ref-CR4" id="ref-link-section-d30854801e565">2010</a>) and then to plan and perform a relatively complex motor task (Vierck <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2006" title="Vierck CJ (2006) Animal models of pain. In: McMahon S, Koltzenburg M (eds) Wall and Melzack’s textbook of pain. Churchill, London" href="/article/10.1007/s10539-014-9469-4#ref-CR123" id="ref-link-section-d30854801e568">2006</a>). Higher-level brain activity (involving the cerebrum) is essential for avoidance learning since decerebrate rats fail to learn to avoid electric shock (Vierck <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2006" title="Vierck CJ (2006) Animal models of pain. In: McMahon S, Koltzenburg M (eds) Wall and Melzack’s textbook of pain. Churchill, London" href="/article/10.1007/s10539-014-9469-4#ref-CR123" id="ref-link-section-d30854801e571">2006</a>). Interestingly, rats exhibit an escape response substantially faster than a brainstem reflex (such as paw licking or jumping) in response to a noxious stimulus (Vierck <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2006" title="Vierck CJ (2006) Animal models of pain. In: McMahon S, Koltzenburg M (eds) Wall and Melzack’s textbook of pain. Churchill, London" href="/article/10.1007/s10539-014-9469-4#ref-CR123" id="ref-link-section-d30854801e574">2006</a>). In addition, the rat threshold for escape response from cold temperatures is approximately 16 °C whereas the threshold for brainstem reflexes is <5 °C (Vierck <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2006" title="Vierck CJ (2006) Animal models of pain. In: McMahon S, Koltzenburg M (eds) Wall and Melzack’s textbook of pain. Churchill, London" href="/article/10.1007/s10539-014-9469-4#ref-CR123" id="ref-link-section-d30854801e577">2006</a>). These comparisons between brainstem reflexes and higher-level escape responses suggest that the cerebrum quickly perceives noxious stimuli as potentially harmful before they are actually physically damaging. Taken together, these results are consistent with rats feeling pain.</p><p>Operant conditioning with negative reinforcement demonstrates that fish can also learn to associate a conditioned stimulus (light cue) with an impending unconditioned stimulus (electric shock) administered in one chamber of a two-chamber holding tank (Hurtado-Parrado <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2010" title="Hurtado-Parrado C (2010) Neuronal mechanisms of learning in teleost fish. Univ Psychol 9:663–672" href="/article/10.1007/s10539-014-9469-4#ref-CR41" id="ref-link-section-d30854801e583">2010</a>). Fish typically learn to terminate their exposure to the electric shock by escaping to the chamber where the shock is not present. With more and more trials, the fish learn to associate the light stimulus with the temporally delayed electric shock and hence begin to escape prior to the delivery of the shock. However, as pointed out above, the escape response in fish is a reflex behaviour and does not equate to the more complex escape routines used in rodent models of pain (Cain et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2010" title="Cain CK, Choi J-S, LeDoux JE (2010) Active avoidance and escape learning. Encycl Behav Neurosci 1:1–9" href="/article/10.1007/s10539-014-9469-4#ref-CR16" id="ref-link-section-d30854801e586">2010</a>). Thus, the better explanation is that fish reflexively associate the stimulus with the shock.</p><p>It has been reasoned that if a behavioural response was modifiable under different circumstances, then it was not a reflex. This vague distinction between reflex and non-reflexive (or flexible) behaviours in fish relies on the notion that higher-level brain activity was associated with the latter and not the former (Dunlop et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2006" title="Dunlop R, Millsopp S, Laming P (2006) Avoidance learning in goldfish (Carassius auratus) and implications for pain perception. Appl Anim Behav Sci 976:255–271" href="/article/10.1007/s10539-014-9469-4#ref-CR27" id="ref-link-section-d30854801e592">2006</a>; Braithwaite et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2013" title="Braithwaite VA, Huntingford F, van den Bos R (2013) Variation in emotion and cognition in fish. J Agric Environ Ethics 26:7–23" href="/article/10.1007/s10539-014-9469-4#ref-CR12" id="ref-link-section-d30854801e595">2013</a>). Evidence for this activity was purported to come from numerous observations that telencephalon ablation perturbed avoidance learning in fish. However, it has been consistently reported that although avoidance learning by fish is perturbed by full or partial forebrain ablations, these animals continue to exhibit escape responses (and many continue to learn to avoid) as a result of electric shock (Hainsworth et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1967" title="Hainsworth FR, Overmier JB, Snowdon CT (1967) Specific and permanent deficits in instrumental avoidance responding following forebrain ablation in the goldfish. J Comp Physiol Psychol 63:111–116" href="/article/10.1007/s10539-014-9469-4#ref-CR39" id="ref-link-section-d30854801e598">1967</a>; Kaplan and Aronson <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1967" title="Kaplan H, Aronson LR (1967) Effect of forebrain ablation on the performance of a conditioned avoidance response in the teleost fish, Tilapia H. Macrocephala. Anim Behav 15:438–448" href="/article/10.1007/s10539-014-9469-4#ref-CR51" id="ref-link-section-d30854801e601">1967</a>; Savage <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1968" title="Savage GE (1968) Temporal factors in avoidance learning in normal and forebrainless goldfish (Cartassius auratus). Nature 218:1168–1169" href="/article/10.1007/s10539-014-9469-4#ref-CR103" id="ref-link-section-d30854801e604">1968</a>; Overmier and Gross <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1974" title="Overmier JB, Gross D (1974) Effects of telencephalic ablation upon nest-building and avoidance bahaviors in East African mouthbreeding fish, Tilapia mossambica. Behav Biol 12:211–222" href="/article/10.1007/s10539-014-9469-4#ref-CR82" id="ref-link-section-d30854801e608">1974</a>; Flood et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1976" title="Flood NC, Overmier JB, Savage GE (1976) Teleost telencephalon and learning: an interpretive review of data and hypotheses. Physiol Behav 16:783–798" href="/article/10.1007/s10539-014-9469-4#ref-CR32" id="ref-link-section-d30854801e611">1976</a>; Overmier and Papini <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1985" title="Overmier JB, Papini MR (1985) Serial ablations of the telencephalon and avoidance learning by goldfish (Carassius auratus). Behav Neurosci 99:509–520" href="/article/10.1007/s10539-014-9469-4#ref-CR83" id="ref-link-section-d30854801e614">1985</a>; Portavella et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2003" title="Portavella M, Salas CM, Vargas JP, Papini MR (2003) Involvement of the telencephalon in spaced-trial avoidance learning in the goldfish (Carassius auratus). Physiol Behav 80:49–56" href="/article/10.1007/s10539-014-9469-4#ref-CR89" id="ref-link-section-d30854801e617">2003</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2004a" title="Portavella M, Torres B, Salas C (2004a) Avoidance response in goldfish: emotional and temporal involvement of medial and lateral telencephalic pallium. J Neurosci 24:2335–2342" href="/article/10.1007/s10539-014-9469-4#ref-CR90" id="ref-link-section-d30854801e620">2004a</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference b" title="Portavella M, Torres B, Salas C, Papini MR (2004b) Lesions of the medial pallium, but not of the lateral pallium, disrupt spaced-trial avoidance learning in goldfish (Carassius auratus). Neurosci Lett 362:75–78" href="/article/10.1007/s10539-014-9469-4#ref-CR91" id="ref-link-section-d30854801e623">b</a>; Portavella and Vargas <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2005" title="Portavella M, Vargas JP (2005) Emotional and spatial learning in goldfish is dependent on different telencephalic pallial systems. Eur J Neurosci 21:2800–2806" href="/article/10.1007/s10539-014-9469-4#ref-CR88" id="ref-link-section-d30854801e627">2005</a>; Vargas et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2009" title="Vargas JP, Lopez JC, Portavella M (2009) What are the functions of fish brain pallium? Brain Res Bull 79:436–440" href="/article/10.1007/s10539-014-9469-4#ref-CR131" id="ref-link-section-d30854801e630">2009</a>). Thus, forebrainless fish are still able to either escape from, or learn (albeit more slowly) to avoid, an electric shock. Fish with, or without, the forebrain had similar latencies of escape. Escape latency was the time taken for a fish to escape from the chamber once it received a shock. Clearly, the forebrain was not needed for fish to exhibit escape behaviour, but it was important for learning the association between the light and the unconditioned stimulus (shock).</p><p>Taken together, the above results demonstrate that the escape responses used in the avoidance learning paradigms for fish involve sub-forebrain regions associated with instinctive and/or reflexive behaviours. Thus, the avoidance learning paradigms typically used in fish studies are more informative about learning processes in fish, then about the sensation of pain experienced by these animals. It is most likely that the sorts of avoidance learning exhibited to date in fish studies is better explained by innate neural circuitry mediating reflex behaviour.</p><h3 class="c-article__sub-heading" id="Sec8">Pain is not essential for reducing injury</h3><p>The idea that nociception has an evolutionary survival advantage for animals is well established in the scientific literature (Kavaliers <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1988" title="Kavaliers M (1988) Evolutionary and comparative aspects of nociception. Brain Res Bull 21:923–931" href="/article/10.1007/s10539-014-9469-4#ref-CR52" id="ref-link-section-d30854801e643">1988</a>). However, the significance of feeling pain in animals is less well understood since the nociception-pain axis has not been carefully interrogated. It has been assumed that pain enables animals to adopt longer-term protective behaviours in order to facilitate tissue repair and to prevent compounding injuries (Bolles and Faneslow <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1980" title="Bolles RC, Faneslow MS (1980) A perceptual-defensive-recuperative model of fear and pain. Behav Brain Sci 3:291–323" href="/article/10.1007/s10539-014-9469-4#ref-CR8" id="ref-link-section-d30854801e646">1980</a>).</p><p>If fish were to feel pain then one would, at least, expect them to exhibit a longer-term protective response to injury. The fins of fish are densely innervated by sensory axons and are one of the most highly sensitive regions of the fish body surface to noxious stimulation (Chervova <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1997" title="Chervova LS (1997) Pain sensitivity of behavior of fishes. J Ichthyol 37:98–102" href="/article/10.1007/s10539-014-9469-4#ref-CR21" id="ref-link-section-d30854801e652">1997</a>). If fish were experiencing pain, and if pain was serving a protective function, then fish should respond to fin injury either by not using that fin or by altering swimming behaviour until the injury was repaired. However, after either partial or complete tail fin amputation, fish show no evidence of protecting their fins by reducing their swimming behaviour; they are instead quite capable of swimming continuously against a current (Fu et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2013" title="Fu C, Cao ZD, Fu SJ (2013) The effects of caudal fin amputation on metabolic interaction between digestion and locomotion in juveniles of three cyprinid fish species with different metabolic modes. Comp Biochem Physiol A: Mol Integr Physiol 164:456–465" href="/article/10.1007/s10539-014-9469-4#ref-CR33" id="ref-link-section-d30854801e655">2013</a>). These observations are also consistent with the normal behaviour of fish with bacterial tail or fin rot. This disease causes progressive erosion of the affected fins/tail and yet these fish swim and eat normally. The consensus in the fish welfare literature is that fin rot, despite its ability to cause loss of most of the tail fin, does not affect the behaviour of fish. These animals continue to eat and swim like their healthy counterparts (Ellis et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2008" title="Ellis T, Oidtmann B, St Hilaire S, Turnbull JF, North BP, MacIntyre CM, Nikolaidis J, Hoyle I, Kestin SC, Knowles TG (2008) Fin erosion in farmed fish. In: Branson EJ (ed) Fish welfare, chapter 9. Wiley, New York. ISBN 978-0-470-69804-4" href="/article/10.1007/s10539-014-9469-4#ref-CR29" id="ref-link-section-d30854801e658">2008</a>). The most plausible interpretation of these observations is that fish do not modulate long-term behaviour in order to allow injury repair. This conclusion is more consistent with fish not feeling pain.</p></div></div></section><section data-title="What is the neural basis of pain?"><div class="c-article-section" id="Sec9-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec9">What is the neural basis of pain?</h2><div class="c-article-section__content" id="Sec9-content"><p>I have suggested above that the behavioural responses of fish to noxious stimuli is best explained by sub-telencephalic reflexes mediated by innate neural circuits rather then by fish experiencing phenomenal consciousness. By accepting this argument it now becomes possible to better address the necessary anatomical prerequisites underlying phenomenal consciousness. All chordates possess a central nervous system consisting of an enlarged anterior end and a posterior cord-like structure. The differences in neuroanatomy that have emerged during evolution within this phylum reflect specialised functions (Butler <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2000" title="Butler AB (2000) Chordate evolution and the origin of the craniates: an old brain in a new head. Anat Rec 261:111–125" href="/article/10.1007/s10539-014-9469-4#ref-CR14" id="ref-link-section-d30854801e670">2000</a>). While the posterior cord has typically preserved a simple morphology that subserves basic locomotor behaviours, the rostral nervous system has instead undergone extensive structural modifications that have led to devise functional consequences. For instance, the evolution of the neocortex in humans has allowed us to experience our environment through subjective mental states such as pain, smell, hearing and vision. By understanding how our environment subjectively “feels” it has become possible for humans to appreciate and predict how other people would respond in certain situations. Consequently by manipulating our environment we are able to affect the behaviour of others to achieve specific outcomes. The human neocortex is particularly adept at this function and it is clearly an important driving force in our cultural evolution.</p><p>What is so unique about the cortex that enables inner mental states? First, the cortex is parcellated into discrete anatomically structures or cortical areas that process information related to specific functions. It is estimated that there are about 200 cortical areas in humans (Kaas <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2012" title="Kaas JH (2012) Evolution of columns, modules, and domains in the neocortex of primates. PNAS 109:10655–10660" href="/article/10.1007/s10539-014-9469-4#ref-CR49" id="ref-link-section-d30854801e676">2012</a>). For instance, the cortical visual system consists of over a dozen distinct regions with diverse subfunctions that are strongly interconnected by reciprocal axon pathways. One of the defining features of these subregions is that they become simultaneously active. Both recurrent activity and binding of neural activity across cortical regions are believed to be essential prerequisites for the subjective experience of vision (Sillito et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2006" title="Sillito AM, Cudeiro J, Jones HE (2006) Always returning: feedback and sensory processing in visual cortex and thalamus. TINS 29:307–316" href="/article/10.1007/s10539-014-9469-4#ref-CR110" id="ref-link-section-d30854801e679">2006</a>; Pollen <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2011" title="Pollen DA (2011) On the emergence of primary visual perception. Cereb Cortex 21:1941–1953" href="/article/10.1007/s10539-014-9469-4#ref-CR87" id="ref-link-section-d30854801e682">2011</a>; Koivisto and Silvanto <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2012" title="Koivisto M, Silvanto J (2012) Visual feature binding: the critical time windows of V1/V2 and parietal activity. Neuroimage 59:1608–1614" href="/article/10.1007/s10539-014-9469-4#ref-CR54" id="ref-link-section-d30854801e685">2012</a>). It has been shown that when neural processing of recurrent signalling from higher cortical regions entering the V1 visual cortex is perturbed by transcranial magnetic stimulation, the subjective awareness of a visual stimulus is disrupted (Koivisto et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2010" title="Koivisto M, Mantyla T, Silvanto J (2010) The role of early visual corex (V1/V2) in conscious and unconscious visual perception. Neuroimage 51:828–834" href="/article/10.1007/s10539-014-9469-4#ref-CR55" id="ref-link-section-d30854801e688">2010</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2011" title="Koivisto M, Railo H, Salminen-Vaparanta N (2011) Transcranial magnetic stimulation of early visual cortex interferes with subjective visual awareness and objective forced-choice performance. Conscious Cogn 20:288–298" href="/article/10.1007/s10539-014-9469-4#ref-CR56" id="ref-link-section-d30854801e692">2011</a>; Jacobs et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2012" title="Jacobs C, de Gaff TA, Goebel R, Sack AT (2012) The temporal dynamics of early visual cortex involvement in behavioral priming. PLoS ONE 7:e48808" href="/article/10.1007/s10539-014-9469-4#ref-CR43" id="ref-link-section-d30854801e695">2012</a>; Railo and Koivisto <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2012" title="Railo H, Koivisto M (2012) Two means of suppressing visual awareness: a direct comparison of visual masking and transcranial magnetic stimulation. Cortex 48:333–343" href="/article/10.1007/s10539-014-9469-4#ref-CR94" id="ref-link-section-d30854801e698">2012</a>; Avanzini et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2013" title="Avanzini P, Fabbri-Destro M, Campi C, Pascarella A, Barchiesi G, Cattaneo L, Rizzolatti G (2013) Spatioptemporal dynamics in understanding hand-object interactions. PNAS 110:15878–15885" href="/article/10.1007/s10539-014-9469-4#ref-CR2" id="ref-link-section-d30854801e701">2013</a>).</p><p>The subregionalisation of the neocortex also allows the formation of spatial maps of the sensory world, such as those associated with the representations of the surface of the body or the visual field. These topographical maps are important for the multiscale processing of sensory information (Kaas <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1997" title="Kaas JH (1997) Topographic maps are fundamental to sensory processing. Brain Res Bull 44:107–112" href="/article/10.1007/s10539-014-9469-4#ref-CR48" id="ref-link-section-d30854801e707">1997</a>; Thivierge and Marcus <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2007" title="Thivierge J-P, Marcus GF (2007) The topographic brain: from neural connectivity to cognition. TINS 30:251–258" href="/article/10.1007/s10539-014-9469-4#ref-CR118" id="ref-link-section-d30854801e710">2007</a>). Variation in the size of the maps alters the sensitivity of responses to stimuli while spatial segregation of neurons responding to selective parts of a stimulus allows for finer perceptual discrimination. Painful and non-painful somatosensory stimuli are topographically mapped to overlying regions in the primary somatosensory cortex (SI) in humans (Mancini et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2012" title="Mancini F, Haggard P, Iannetti GD, Longo MR, Sereno MI (2012) Fine-grained nociceptive maps in primary somatosensory cortex. J Neurosci 32:17155–17162" href="/article/10.1007/s10539-014-9469-4#ref-CR63" id="ref-link-section-d30854801e713">2012</a>). These results are consistent with the known point-to-point topography from the body surface to SI (called somatotopy) that underlies spatial acuity. However, by using high resolution mapping in the squirrel monkey SI (sub-millimetre level) it was revealed that there were slight differences in the localisation of different somatosensory modalities (Chen et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2001" title="Chen LM, Friedman RM, Ramsden BM, LaMotte RH, Roe AW (2001) Fine-scale organization of SI (area 3b) in the squirrel monkey revealed with intrinsic optical imaging. J Neurophysiol 86:3011–3029" href="/article/10.1007/s10539-014-9469-4#ref-CR19" id="ref-link-section-d30854801e716">2001</a>). This slight physical separation of cortical neurons responding to different peripheral stimuli suggests that differences in the subjective quality of somatosensory sensations may arise as early as in SI. Somatotopic maps for painful stimuli are also present in the human SII and insular cortices (Baumgartner et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2010" title="Baumgartner U, Iannetti GD, Zambreanu L, Stoeter P, Treede R-D, Tracey I (2010) Multiple somatotopic representations of heat and mechanical pain in the operculo-insular cortex: a high-resolution fMRI study. J Neurophysiol 104:2863–2872" href="/article/10.1007/s10539-014-9469-4#ref-CR150" id="ref-link-section-d30854801e719">2010</a>). Interestingly, different qualities of painful stimuli (such as heat and pinprick) are more distinctly mapped topographically to different regions of SII and the insular cortex than in SI. Similarly, painful and non-painful stimuli are mapped to separate regions in human SII (Torquati et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2005" title="Torquati K, Pizzella V, Babiloni C, del Gratta C, Della Penna S, Ferretti A, Franciotti R, Rossini PM, Romani GL (2005) Nociceptive and non-nociceptive sub-regions in the human somatosensory cortex: an MEG study using fMRI constraints. Neuroimage 26:48–56" href="/article/10.1007/s10539-014-9469-4#ref-CR120" id="ref-link-section-d30854801e723">2005</a>). This separation of cortical processing of heat and tactile stimuli within different cortical areas has also been observed in non-human primates (Chen et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2011" title="Chen LM, Dillenburger BC, Wang F, Friedman RM, Avison MJ (2011) High-resolution functional magnetic resonance imaging of noxious heat and activations along the central sulcus in New World monkeys. Pain 152:522–532" href="/article/10.1007/s10539-014-9469-4#ref-CR20" id="ref-link-section-d30854801e726">2011</a>). These multiple neural maps suggests that SII and the insular cortex play important roles in discriminating differences in the subjective quality of somatosensory stimuli, particularly painful from non-painful (Tommerdahl et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1996" title="Tommerdahl M, Delemos KA, Vierck CJ, Favorov OV, Whitsel BL (1996) Anterior parietal cortical response to tactile and skin-heating stimuli applied to the same skin site. J Neurophysiol 75:2662–2670" href="/article/10.1007/s10539-014-9469-4#ref-CR119" id="ref-link-section-d30854801e729">1996</a>; Baumgartner et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2010" title="Baumgartner U, Iannetti GD, Zambreanu L, Stoeter P, Treede R-D, Tracey I (2010) Multiple somatotopic representations of heat and mechanical pain in the operculo-insular cortex: a high-resolution fMRI study. J Neurophysiol 104:2863–2872" href="/article/10.1007/s10539-014-9469-4#ref-CR150" id="ref-link-section-d30854801e732">2010</a>; Chen et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2011" title="Chen LM, Dillenburger BC, Wang F, Friedman RM, Avison MJ (2011) High-resolution functional magnetic resonance imaging of noxious heat and activations along the central sulcus in New World monkeys. Pain 152:522–532" href="/article/10.1007/s10539-014-9469-4#ref-CR20" id="ref-link-section-d30854801e735">2011</a>; Mazzola et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2012" title="Mazzola L, Faillenot I, Barral F-G, Mauguiere F, Peyron R (2012) Spatial segregation of somatosensory and pain activations in the human operculo-insular cortex. Neuroimage 60:5409–5418" href="/article/10.1007/s10539-014-9469-4#ref-CR68" id="ref-link-section-d30854801e738">2012</a>). This idea is supported by evidence from direct electrical stimulation of discrete areas in the human insular cortex (Afif et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2010" title="Afif A, Minotti L, Kahane P, Hoffman D (2010) Anatomofunctional organization of the insular cortex: a study using intracerebral electrical stimulation in epileptic patients. Epilepsia 51:2305–2315" href="/article/10.1007/s10539-014-9469-4#ref-CR1" id="ref-link-section-d30854801e742">2010</a>).</p><p>Second, the cortex is a laminated structure that enables the efficient processing and integration of different types of neural information by unique subpopulations of neurons (Schubert et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2007" title="Schubert D, Kotter R, Staiger JF (2007) Mapping functional connectivity in barrel-related columns reveals layer- and cell type-specific microcircuits. Brain Struct Funct 212:107–119" href="/article/10.1007/s10539-014-9469-4#ref-CR105" id="ref-link-section-d30854801e748">2007</a>; Maier et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2010" title="Maier A, Adams GK, Aura C, Leopold DA (2010) Distinct superficial and deep laminar domains of activity in the visual cortex during rest and stimulation. Front Syst Neurosci. doi:
 10.3389/fnsys.2010.00031
 
 
 " href="/article/10.1007/s10539-014-9469-4#ref-CR61" id="ref-link-section-d30854801e751">2010</a>; Larkum <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2013" title="Larkum M (2013) A cellular mechanism for cortical associations: an organizing principle for the cerebral cortex. TINS 36:141–151" href="/article/10.1007/s10539-014-9469-4#ref-CR59" id="ref-link-section-d30854801e754">2013</a>). Lamination appears to facilitate complex wiring patterns during development. If two populations of neurons were randomly distributed within a specific brain region and incoming axons were required to synapse with only one subpopulation, then those axons would need to rely on stochastic and hence error-prone searching to complete wiring. On the other hand, when similar neurons are partitioned together in a single lamina then a small set of molecular cues is able to guide axons with high precision to their appropriate post-synaptic target. Two principal afferent inputs (from the neocortex itself, and the thalamus) enter the neocortex and separately innervate distinct layers (Nieuwenhuys <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1994" title="Nieuwenhuys R (1994) The neocortex. An overview of its evolutionary development, structural organization and synaptology. Anat Embryol 190:307–337" href="/article/10.1007/s10539-014-9469-4#ref-CR74" id="ref-link-section-d30854801e757">1994</a>). The main thalamic fibres terminate densely in layer IV (called the granular layer) while the neocortical fibres innervate different pyramidal neurons in layers I–III (supragranular layers) (Opris <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2013" title="Opris I (2013) Inter-laminar microcircuits across neocortex: repair and augmentation. Front Syst Neurosci 7:80" href="/article/10.1007/s10539-014-9469-4#ref-CR80" id="ref-link-section-d30854801e760">2013</a>). By selectively ablating Pax6, a developmentally significant patterning gene, in the cortex of mice it is possible to disrupt the laminar organisation of this structure (Tuoc et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2009" title="Tuoc TC, Radyushkin K, Tonchev AB, Piñon MC, Ashery-Padan R, Molnár Z, Davidoff MS, Stoykova A (2009) Selective cortical layering abnormalities and behavioral deficits in cortex-specific Pax6 knock-out mice. J Neurosci 29:8335–8349" href="/article/10.1007/s10539-014-9469-4#ref-CR121" id="ref-link-section-d30854801e764">2009</a>). This altered cortical layering causes neurological deficits that are similar to those observed in humans with Pax6 haploinsufficiency (Tuoc et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2009" title="Tuoc TC, Radyushkin K, Tonchev AB, Piñon MC, Ashery-Padan R, Molnár Z, Davidoff MS, Stoykova A (2009) Selective cortical layering abnormalities and behavioral deficits in cortex-specific Pax6 knock-out mice. J Neurosci 29:8335–8349" href="/article/10.1007/s10539-014-9469-4#ref-CR121" id="ref-link-section-d30854801e767">2009</a>) and provides strong experimental evidence of the importance of lamination to cortical function. A number of human brain disorders involve defects in cortical lamination that are detrimental to brain function (Guerrini et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2008" title="Guerrini R, Dobyns WB, Barkovich AJ (2008) Abnormal development of the human cerebral cortex: genetics, functional consequences and treatment options. TINS 31:154–162" href="/article/10.1007/s10539-014-9469-4#ref-CR36" id="ref-link-section-d30854801e770">2008</a>; Guerrini and Parrini <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2010" title="Guerrini R, Parrini E (2010) Neuronal migration disorders. Neurobiol Dis 38:154–166" href="/article/10.1007/s10539-014-9469-4#ref-CR35" id="ref-link-section-d30854801e773">2010</a>; Bozzi et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2012" title="Bozzi Y, Casarosa S, Caleo M (2012) Epilepsy as a neurodevelopmental disorder. Front Psychol. doi:
 10.3389/fpsyt.2012.00019
 
 
 " href="/article/10.1007/s10539-014-9469-4#ref-CR9" id="ref-link-section-d30854801e776">2012</a>).</p><p>Third, lamination facilitates the economical establishment of microcircuitry between neurons processing different properties of the stimulus. A vertical canonical microcircuit is established which leads to the emergence of functionally interconnected columns and minicolumns of neurons (Mountcastle <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1997" title="Mountcastle VB (1997) The columnar organization of the neocortex. Brain 120:701–722" href="/article/10.1007/s10539-014-9469-4#ref-CR70" id="ref-link-section-d30854801e783">1997</a>). For example, a hexagonal column in the primate somatosensory cortex is about 400 μm in width and contains populations of neurons that respond to the same stimulus (e.g. light touch or joint stimulation) arising from a specific topographical zone of the body. Columns can be associated with processing information related to a specific function (e.g. “visual tracking” and “arm reach” columns in the parietal cortex; Kass <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2012" title="Kaas JH (2012) Evolution of columns, modules, and domains in the neocortex of primates. PNAS 109:10655–10660" href="/article/10.1007/s10539-014-9469-4#ref-CR49" id="ref-link-section-d30854801e786">2012</a>). Each column itself consists of minicolumns (80–100 neurons) that are ~30–50 μm in diameter and interconnected by short-range horizontal processes (Buxhoeveden and Casanova <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2002" title="Buxhoeveden DP, Casanova MF (2002) The minicolumn hypothesis in neuroscience. Brain 125:935–951" href="/article/10.1007/s10539-014-9469-4#ref-CR15" id="ref-link-section-d30854801e789">2002</a>). While columns are most clearly distinguished in the sensory and motor cortices of primates, minicolumns appear to be ubiquitous in all animals with a neocortex (Kaas <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2012" title="Kaas JH (2012) Evolution of columns, modules, and domains in the neocortex of primates. PNAS 109:10655–10660" href="/article/10.1007/s10539-014-9469-4#ref-CR49" id="ref-link-section-d30854801e792">2012</a>). Minicolumns have a small receptive field within the larger receptive field of the column. The correlated activity in the fine-scale networks of minicolumns produces concentrated bursts of neural activity that may enable the cortex to transmit signals in the face of background noise (Ohiorhenuan et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2010" title="Ohiorhenuan IE, Mechler F, Purpura KP, Schmid AM, Hu Q, Victor JD (2010) Sparse coding and high-order correlations in fine-scale cortical networks. Nature 466:617–622" href="/article/10.1007/s10539-014-9469-4#ref-CR79" id="ref-link-section-d30854801e795">2010</a>). The function of the cortex seems to depend on the ability of canonical circuitry within the minicolumns to rapidly switch from feedforward to feedback processing between layers. During learned tasks in responses to cues in the awake monkey, information flows from layer 4 to layer 2/3 and then down to layer 5 in a feedforward loop in the temporal neocortex (Takeuchi et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2011" title="Takeuchi D, Hirabayashi T, Tamura K, Miyashita Y (2011) Reversal of interlaminar signal between sensory and memory processing in monkey temporal cortex. Science 331:1443–1447" href="/article/10.1007/s10539-014-9469-4#ref-CR116" id="ref-link-section-d30854801e799">2011</a>; Bastos et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2012" title="Bastos AM, Usrey WM, Adams RA, Mangun GR, Fries P, Friston KJ (2012) Canonical microcircuits for predictive coding. Neuron 76:695–711" href="/article/10.1007/s10539-014-9469-4#ref-CR5" id="ref-link-section-d30854801e802">2012</a>). This is followed shortly afterwards by a feedback loop from layer 5 to layer 2/3. Correlated firing of layer 2/3 and layer 5 neurons in minicolumns occurs during decision making in the monkey prefrontal cortex, an area responsible for executive control in primates (Opris et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2012" title="Opris I, Fuqua JL, Huetti PF, Gerhardt GA, Berger TW, Hampson RE, Deadwyler SA (2012) Closing the loop in primate prefrontal cortex: inter-laminar processing. Front Neural Circuits 6:1–13" href="/article/10.1007/s10539-014-9469-4#ref-CR81" id="ref-link-section-d30854801e805">2012</a>). The accuracy of error-prone tasks was increased when layer 5 neurons were artificially stimulated by activity recorded during successful task execution. These results provide evidence for the role of the minicolumn as the fundamental processing unit of the neocortex associated with higher order behaviour (Bastos et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2012" title="Bastos AM, Usrey WM, Adams RA, Mangun GR, Fries P, Friston KJ (2012) Canonical microcircuits for predictive coding. Neuron 76:695–711" href="/article/10.1007/s10539-014-9469-4#ref-CR5" id="ref-link-section-d30854801e808">2012</a>; Opris et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2012" title="Opris I, Fuqua JL, Huetti PF, Gerhardt GA, Berger TW, Hampson RE, Deadwyler SA (2012) Closing the loop in primate prefrontal cortex: inter-laminar processing. Front Neural Circuits 6:1–13" href="/article/10.1007/s10539-014-9469-4#ref-CR81" id="ref-link-section-d30854801e811">2012</a>).</p><p>In summary, the unique morphology of the mammalian cortex facilitates multiscale processing of sensory information. Initially there is course scaling at the level of gross anatomical cortical regions specialising, for example, in processing of visual or somatosensory information. Some of these regions are then topographically mapped in order to preserve spatial relationships and facilitate selective processing of specific sensory features. Importantly, to preserve the holistic quality of a sensory stimulus, these subregions are strongly interconnected via axon pathways that create synchronized re-entrant loops of neural activity. Cortical regions are laminated which supports finer scale sensitivity in the processing of specific features. Finally, canonical microcircuits (minicolumns) bridge across layers to enhance signal contrast (Casanova <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2010" title="Casanova MF (2010) Cortical organization: anatomical findings based on systems theory. Transl Neurosci 1:62–71" href="/article/10.1007/s10539-014-9469-4#ref-CR17" id="ref-link-section-d30854801e817">2010</a>). Local connectivity between minicolumns enables the lowest level of stimulus binding that contributes to the holistic nature of the stimulus (Buxhoeveden and Casanova <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2002" title="Buxhoeveden DP, Casanova MF (2002) The minicolumn hypothesis in neuroscience. Brain 125:935–951" href="/article/10.1007/s10539-014-9469-4#ref-CR15" id="ref-link-section-d30854801e820">2002</a>).</p><p>I propose that only animals possessing the above neuroanatomical features (i.e. discrete cortical sensory regions, topographical maps, multiple cortical layers, columns/minicolumns and strong local and long-range interconnections), or their functionally analogous counterparts, have the necessary morphological prerequisites for experiencing subjective inner mental states such as pain. It has been argued that since the avian pallium is non-laminated, and yet these animals exhibit high levels of cognitive ability and behaviours rivalling those of primates, that lamination is not an essential prerequisite for consciousness (Gunturkun <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2005" title="Gunturkun O (2005) The avian ‘prefrontal cortex’ and cognition. Curr Opin Neurobiol 15:686–693" href="/article/10.1007/s10539-014-9469-4#ref-CR37" id="ref-link-section-d30854801e826">2005</a>; Kirsch et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2008" title="Kirsch JA, Gunturkun O, Rose J (2008) Insight without cortex: lessons from the avian brain. Conscious Cogn 17:475–483" href="/article/10.1007/s10539-014-9469-4#ref-CR53" id="ref-link-section-d30854801e829">2008</a>; Gunturkun <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2012" title="Gunturkun O (2012) The convergent evolution of neural substrates for cognition. Psychol Res 76:212–219" href="/article/10.1007/s10539-014-9469-4#ref-CR38" id="ref-link-section-d30854801e832">2012</a>; Veit and Nieder <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2013" title="Veit L, Nieder A (2013) Abstract rule neurons in the endbrain support intelligent behaviour in corvid songbirds. Nat Commun 4:2878. doi:
 10.1038/ncomms3878
 
 
 " href="/article/10.1007/s10539-014-9469-4#ref-CR122" id="ref-link-section-d30854801e835">2013</a>). However, the classic view of the organisation of the avian telencephalon has been revised and previous subpallial regions are now recognised as pallial in nature (Shimizu <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2009" title="Shimizu T (2009) Why can birds be so smart? Background, significance, and implications of the revised view of the avian brain. Comp Cogn Behav Rev 4:103–115" href="/article/10.1007/s10539-014-9469-4#ref-CR107" id="ref-link-section-d30854801e838">2009</a>). Careful examination of pallial neuroanatomy has further revealed that distinct regions of the avian pallium act like layers of the neocortex (Dugas-Ford et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2012" title="Dugas-Ford J, Rowell JJ, Ragsdale CW (2012) Cell-type homologies and the origins of the neocortex. PNAS 109:16974–16979" href="/article/10.1007/s10539-014-9469-4#ref-CR25" id="ref-link-section-d30854801e842">2012</a>). Moreover, columnar processing units appear to operate across these brain regions in the processing of sensory and motor information (Jarvis et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2013" title="Jarvis ED, Yu J, Rivas MV, Horita H, Feenders G, Whitney O, Jarvis SC, Jarvis ER, Kubikova L, Puck AEP, Siang-Bakshi C, Martin S, McElroy M, Hara E, Howard J, Pfenning A, Mouritsen H, Chen C-C, Wada K (2013) Global view of the functional molecular organization of the avian cerebrum: mirror images and functional columns. J Comp Neurol 521:3614–3665" href="/article/10.1007/s10539-014-9469-4#ref-CR44" id="ref-link-section-d30854801e845">2013</a>). When this is combined with complex parcellation, the presence of topographical maps and strong interconnectivity in the avian pallium (Shimizu et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1995" title="Shimizu T, Cox K, Karten HJ (1995) Intratelencephalic projections of the visual wulst in pigeons (Columba livia). J Comp Neurol 359:551–572" href="/article/10.1007/s10539-014-9469-4#ref-CR109" id="ref-link-section-d30854801e848">1995</a>; Shimizu and Bowers <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1999" title="Shimizu T, Bowers AN (1999) Visual circuits of the avian telencephalon. Behav Brain Res 98:183–191" href="/article/10.1007/s10539-014-9469-4#ref-CR108" id="ref-link-section-d30854801e851">1999</a>; Bingman and Able <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2002" title="Bingman VP, Able KP (2002) Maps in birds: representational mechanisms and neural bases. Curr Opin Neurobiol 12:745–750" href="/article/10.1007/s10539-014-9469-4#ref-CR7" id="ref-link-section-d30854801e854">2002</a>; Manger et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2002" title="Manger PR, Elston GN, Pettigrew JD (2002) Multiple maps and activity-dependent representational plasticity in the anterior Wust of the adult barn owl (Tyto alba). Eur J Neurosci 16:743–750" href="/article/10.1007/s10539-014-9469-4#ref-CR65" id="ref-link-section-d30854801e857">2002</a>; Nguyen et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2004" title="Nguyen AP, Spetch ML, Crowder NA, Winship IR, Hurd PL, Wylie DR (2004) A dissociation of motion and spatial-pattern vision in the avian telencephalon: implications for the evolution of “visual streams”. J Neurosci 24:4962–4970" href="/article/10.1007/s10539-014-9469-4#ref-CR73" id="ref-link-section-d30854801e861">2004</a>; Watanabe and Masuda <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2010" title="Watanabe S, Masuda S (2010) Integration of auditory and visual information in human face discrimination in pigeons. Behavioral and anatomical study. Behav Brain Res 207:61–69" href="/article/10.1007/s10539-014-9469-4#ref-CR124" id="ref-link-section-d30854801e864">2010</a>), it appears that birds possess the necessary neural machinery for phenomenal consciousness.</p><p>The pallium of fish is non-laminated. It is partitioned into five broad nuclear regions (dorsomedial, dorsolateral, dorsodorsal, dorsoposterior and ventral; Northcutt <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2011" title="Northcutt RG (2011) Do teleost fishes possess a homolog of mammalian isocortex? Brain Behav Evol 78:136–138" href="/article/10.1007/s10539-014-9469-4#ref-CR77" id="ref-link-section-d30854801e870">2011</a>). While the dorsodorsal pallium is believed to be homologous to the neocortex there remains some controversy as to the definitive homology between these structures (Echleter and Saidel <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1981" title="Echleter SM, Saidel WM (1981) Forebrain connections in the goldfish support telencephalic homologies with land vertebrates. Science 212:683–685" href="/article/10.1007/s10539-014-9469-4#ref-CR28" id="ref-link-section-d30854801e873">1981</a>; Northcutt <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2008" title="Northcutt RG (2008) Forebrain evolution in bony fishes. Brain Res Bull 75:191–205" href="/article/10.1007/s10539-014-9469-4#ref-CR76" id="ref-link-section-d30854801e876">2008</a>; Braford <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2009" title="Braford MR (2009) Stalking the everted telencephalon: comparisons of forebrain organization in basal ray-finned fishes and teleosts. Brain Behav Evol 74:56–76" href="/article/10.1007/s10539-014-9469-4#ref-CR10" id="ref-link-section-d30854801e879">2009</a>; Northcutt <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2011" title="Northcutt RG (2011) Do teleost fishes possess a homolog of mammalian isocortex? Brain Behav Evol 78:136–138" href="/article/10.1007/s10539-014-9469-4#ref-CR77" id="ref-link-section-d30854801e882">2011</a>). There is converging evidence from electrophysiological recordings (Precht et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1998" title="Precht JC, von der Emde G, Wolfart J, Karamursel S, Akoev GN, Andrianov YN, Bullock TH (1998) Sensory processing in the pallium of a Mormyrid fish. J Neurosci 18:7381–7393" href="/article/10.1007/s10539-014-9469-4#ref-CR92" id="ref-link-section-d30854801e886">1998</a>; Saidel et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2001" title="Saidel WM, Marquez-Houston K, Butler AB (2001) Identification of visual pallial telencephalon in the goldfish, Carassius auratus: a combined cytochrome oxidase and electrophysiological study. Brain Res 919:82–93" href="/article/10.1007/s10539-014-9469-4#ref-CR101" id="ref-link-section-d30854801e889">2001</a>; Northcutt et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2004" title="Northcutt RG, Plassman W, Holmes PH, Saidel WM (2004) A pallial visual area in the telencephalon of the bony fish Polypterus. Brain Behav Evol 64:1–10" href="/article/10.1007/s10539-014-9469-4#ref-CR78" id="ref-link-section-d30854801e892">2004</a>) and neuroanatomical tracing (Yamamoto and Ito <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2008" title="Yamamoto N, Ito H (2008) Visual, lateral line, and auditory ascending pathways to the dorsal telencephalic area through the rostrolateral region of the lateral preglomerular nucleus in Cyprinids. J Comp Neurol 508:615–647" href="/article/10.1007/s10539-014-9469-4#ref-CR129" id="ref-link-section-d30854801e895">2008</a>) that, unlike in the neocortex, sensory information such as visual input, is diffusely processed across the fish dorsal pallium, and certainly not localised to multiple interconnected areas that are topographically mapped (Giassi et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2012" title="Giassi ACC, Ellis W, Maler L (2012) Organization of the gymnotiform fish pallium in relation to learning and memory: III. Intrinsic connections. J Comp Neurol 520:3369–3394" href="/article/10.1007/s10539-014-9469-4#ref-CR34" id="ref-link-section-d30854801e898">2012</a>). Evidence is also lacking for canonical microcircuitry subserving fine scale processing of sensory information in the dorsal pallium. This lack of contrast in signal processing does not support the ability of the fish pallium to differentiate sensory modalities with sufficient resolution to allow the emergence of distinct feelings for different sensory modalities.</p><p>It has been suggested that sub-forebrain structures in fish may somehow take over the function of phenomenal consciousness in the neocortex. While parcellated sensory processing, laminated cytoarchitecture and columnar-like modules are present in the mid- and hindbrains of some fish (Meek <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1983" title="Meek J (1983) Functional anatomy of the tectum mesencephala of the goldfish. An explorative analysis of the functional implications of the laminar structural organization of the tectum. Brain Res 287:247–297" href="/article/10.1007/s10539-014-9469-4#ref-CR69" id="ref-link-section-d30854801e904">1983</a>; Krahe and Maler <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2014" title="Krahe R, Maler L (2014) Neural maps in the electrosensory system of weakly electric fish. Curr Opin Neurobiol 24:13–21" href="/article/10.1007/s10539-014-9469-4#ref-CR57" id="ref-link-section-d30854801e907">2014</a>), these structures lack the necessary local and long-range feedforward and recurrent pathways associated with information binding underlying phenomenal consciousness (Baars et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2013" title="Baars BJ, Franklin S, Ramsoy TZ (2013) Global workspace dynamics: cortical “binding and propagation” enables conscious contents. Front Psychol. doi:
 10.3389/fpsyg.2013.00200
 
 
 " href="/article/10.1007/s10539-014-9469-4#ref-CR3" id="ref-link-section-d30854801e910">2013</a>). Instead, the vertebrate midbrain optic tectum has conserved structural features across a variety of species such as fish, frogs, birds and mammals that subserve common functionalities (e.g. orienting, direction-sensitivity, and spatial relationships; Ingle <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1973" title="Ingle D (1973) Evolutionary perspectives on the function of the optic tectum. Brain Behav Evol 8:211–237" href="/article/10.1007/s10539-014-9469-4#ref-CR42" id="ref-link-section-d30854801e913">1973</a>). Furthermore, while ablation of the tecta perturbs visual function, startle responses in tectumless fish are preserved (Yager et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1977" title="Yager D, Sharma SC, Grover BG (1977) Visual function in goldfish with unilateral and bilateral tectal ablation. Brain Res 137:267–275" href="/article/10.1007/s10539-014-9469-4#ref-CR127" id="ref-link-section-d30854801e916">1977</a>; Roeser and Naier <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2003" title="Roeser T, Naier H (2003) Visuomotor behaviors in larval zebrafish after GFP-guided laser ablation of the optic tectum. J Neurosci 23:3726–3734" href="/article/10.1007/s10539-014-9469-4#ref-CR96" id="ref-link-section-d30854801e920">2003</a>). Thus, the tectum is not needed to respond to somatosensory stimuli and certainly does not possess novel circuitry responsible for pain. On the basis of our current understanding of the structure and function of the “fish” brain, it most likely that fish do not have the necessary neural machinery for phenomenal consciousness.</p><p>In summary, I have demonstrated how misleading it is to infer that fish have feelings on the basis of behavioural responses to sensory stimulation. It is essential that our anthropomorphic tendencies to bestow animals with feelings does not hinder the progress of scientific enquiry into the evolution of phenomenal consciousness. I propose that there are a number of fundamental neural building blocks that are necessary prerequisites for phenomenal consciousness in the vertebrate lineage. The possession of this hardware sets the minimal requirements for the sensation of noxious stimuli as painful. The idea that other neural architectures that have been specifically wired for fundamentally different functions in vertebrates (such as the mid- and hindbrains) could also subserve pain in fish is incongruent with evolutionary biology and neuroscience. While there is some degree of plasticity of function in the mammalian neocortex (Kupers and Pitto <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2013" title="Kupers R, Pitto M (2013) Compensatory plasticity and cross-modal reorganization following early visual deprivation. Neurosci Biobehav Rev. doi:
 10.1016/j.neubiorev.2013.08.001
 
 
 " href="/article/10.1007/s10539-014-9469-4#ref-CR58" id="ref-link-section-d30854801e926">2013</a>), the very notion that either the fish tectum as well as the mid- and hindbrain reticular formations (that are reciprocally interconnected with the tectum; Perez-Perez et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2003" title="Perez-Perez MP, Luque MA, Herrero L, Nunez-Abades PA, Torres B (2003) Connectivity of the goldfish optic tectum with the mesencephalic and rhombencephalic reticular formation. Exp Brain Res 151:123–135" href="/article/10.1007/s10539-014-9469-4#ref-CR84" id="ref-link-section-d30854801e929">2003</a>; Luque et al. <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2005" title="Luque MA, Perez-Perez MP, Herrero L, Torres B (2005) Involvement of the optic tectum and mesencephalic reticular formation in the generation of saccadic eye movements in goldfish. Brain Res Rev 49:388–397" href="/article/10.1007/s10539-014-9469-4#ref-CR60" id="ref-link-section-d30854801e932">2005</a>) has some hidden neural circuitry that allows for the processing of somatosensory inputs into discrete feelings of pinprick, heat, cold, scratch, cutting and stabbing is difficult to defend.</p></div></div></section> </div> <div id="MagazineFulltextArticleBodySuffix"><section aria-labelledby="Bib1" data-title="References"><div class="c-article-section" id="Bib1-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Bib1">References</h2><div class="c-article-section__content" id="Bib1-content"><div data-container-section="references"><ul class="c-article-references" data-track-component="outbound reference" data-track-context="references section"><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR1">Afif A, Minotti L, Kahane P, Hoffman D (2010) Anatomofunctional organization of the insular cortex: a study using intracerebral electrical stimulation in epileptic patients. Epilepsia 51:2305–2315</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 1" href="http://scholar.google.com/scholar_lookup?&title=Anatomofunctional%20organization%20of%20the%20insular%20cortex%3A%20a%20study%20using%20intracerebral%20electrical%20stimulation%20in%20epileptic%20patients&journal=Epilepsia&volume=51&pages=2305-2315&publication_year=2010&author=Afif%2CA&author=Minotti%2CL&author=Kahane%2CP&author=Hoffman%2CD"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR2">Avanzini P, Fabbri-Destro M, Campi C, Pascarella A, Barchiesi G, Cattaneo L, Rizzolatti G (2013) Spatioptemporal dynamics in understanding hand-object interactions. PNAS 110:15878–15885</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 2" href="http://scholar.google.com/scholar_lookup?&title=Spatioptemporal%20dynamics%20in%20understanding%20hand-object%20interactions&journal=PNAS&volume=110&pages=15878-15885&publication_year=2013&author=Avanzini%2CP&author=Fabbri-Destro%2CM&author=Campi%2CC&author=Pascarella%2CA&author=Barchiesi%2CG&author=Cattaneo%2CL&author=Rizzolatti%2CG"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR3">Baars BJ, Franklin S, Ramsoy TZ (2013) Global workspace dynamics: cortical “binding and propagation” enables conscious contents. Front Psychol. doi:<a href="https://doi.org/10.3389/fpsyg.2013.00200" data-track="click_references" data-track-action="external reference" data-track-value="external reference" data-track-label="10.3389/fpsyg.2013.00200">10.3389/fpsyg.2013.00200</a> </p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 3" href="http://scholar.google.com/scholar_lookup?&title=Global%20workspace%20dynamics%3A%20cortical%20%E2%80%9Cbinding%20and%20propagation%E2%80%9D%20enables%20conscious%20contents&journal=Front%20Psychol&doi=10.3389%2Ffpsyg.2013.00200&publication_year=2013&author=Baars%2CBJ&author=Franklin%2CS&author=Ramsoy%2CTZ"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR4">Baastrup C, Maersk-Moller CC, Nyengaard JR, Jensen TS, Finnerup NB (2010) Spinal-, brainstem- and cerebrally mediated responses at- and below-level of a spinal cord contusion in rats: evaluation of pain-like behavior. Pain 151:670–679</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 4" href="http://scholar.google.com/scholar_lookup?&title=Spinal-%2C%20brainstem-%20and%20cerebrally%20mediated%20responses%20at-%20and%20below-level%20of%20a%20spinal%20cord%20contusion%20in%20rats%3A%20evaluation%20of%20pain-like%20behavior&journal=Pain&volume=151&pages=670-679&publication_year=2010&author=Baastrup%2CC&author=Maersk-Moller%2CCC&author=Nyengaard%2CJR&author=Jensen%2CTS&author=Finnerup%2CNB"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR5">Bastos AM, Usrey WM, Adams RA, Mangun GR, Fries P, Friston KJ (2012) Canonical microcircuits for predictive coding. Neuron 76:695–711</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 5" href="http://scholar.google.com/scholar_lookup?&title=Canonical%20microcircuits%20for%20predictive%20coding&journal=Neuron&volume=76&pages=695-711&publication_year=2012&author=Bastos%2CAM&author=Usrey%2CWM&author=Adams%2CRA&author=Mangun%2CGR&author=Fries%2CP&author=Friston%2CKJ"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR150">Baumgartner U, Iannetti GD, Zambreanu L, Stoeter P, Treede R-D, Tracey I (2010) Multiple somatotopic representations of heat and mechanical pain in the operculo-insular cortex: a high-resolution fMRI study. J Neurophysiol 104:2863–2872</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 6" href="http://scholar.google.com/scholar_lookup?&title=Multiple%20somatotopic%20representations%20of%20heat%20and%20mechanical%20pain%20in%20the%20operculo-insular%20cortex%3A%20a%20high-resolution%20fMRI%20study&journal=J%20Neurophysiol&volume=104&pages=2863-2872&publication_year=2010&author=Baumgartner%2CU&author=Iannetti%2CGD&author=Zambreanu%2CL&author=Stoeter%2CP&author=Treede%2CR-D&author=Tracey%2CI"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR7">Bingman VP, Able KP (2002) Maps in birds: representational mechanisms and neural bases. Curr Opin Neurobiol 12:745–750</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 7" href="http://scholar.google.com/scholar_lookup?&title=Maps%20in%20birds%3A%20representational%20mechanisms%20and%20neural%20bases&journal=Curr%20Opin%20Neurobiol&volume=12&pages=745-750&publication_year=2002&author=Bingman%2CVP&author=Able%2CKP"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR8">Bolles RC, Faneslow MS (1980) A perceptual-defensive-recuperative model of fear and pain. Behav Brain Sci 3:291–323</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 8" href="http://scholar.google.com/scholar_lookup?&title=A%20perceptual-defensive-recuperative%20model%20of%20fear%20and%20pain&journal=Behav%20Brain%20Sci&volume=3&pages=291-323&publication_year=1980&author=Bolles%2CRC&author=Faneslow%2CMS"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR9">Bozzi Y, Casarosa S, Caleo M (2012) Epilepsy as a neurodevelopmental disorder. Front Psychol. doi:<a href="https://doi.org/10.3389/fpsyt.2012.00019" data-track="click_references" data-track-action="external reference" data-track-value="external reference" data-track-label="10.3389/fpsyt.2012.00019">10.3389/fpsyt.2012.00019</a> </p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 9" href="http://scholar.google.com/scholar_lookup?&title=Epilepsy%20as%20a%20neurodevelopmental%20disorder&journal=Front%20Psychol&doi=10.3389%2Ffpsyt.2012.00019&publication_year=2012&author=Bozzi%2CY&author=Casarosa%2CS&author=Caleo%2CM"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR10">Braford MR (2009) Stalking the everted telencephalon: comparisons of forebrain organization in basal ray-finned fishes and teleosts. Brain Behav Evol 74:56–76</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 10" href="http://scholar.google.com/scholar_lookup?&title=Stalking%20the%20everted%20telencephalon%3A%20comparisons%20of%20forebrain%20organization%20in%20basal%20ray-finned%20fishes%20and%20teleosts&journal=Brain%20Behav%20Evol&volume=74&pages=56-76&publication_year=2009&author=Braford%2CMR"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR11">Braithwaite VA, Boulcott P (2007) Pain perception, aversion and fear in fish. Dis Aquat Org 75:131–138</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 11" href="http://scholar.google.com/scholar_lookup?&title=Pain%20perception%2C%20aversion%20and%20fear%20in%20fish&journal=Dis%20Aquat%20Org&volume=75&pages=131-138&publication_year=2007&author=Braithwaite%2CVA&author=Boulcott%2CP"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR12">Braithwaite VA, Huntingford F, van den Bos R (2013) Variation in emotion and cognition in fish. J Agric Environ Ethics 26:7–23</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 12" href="http://scholar.google.com/scholar_lookup?&title=Variation%20in%20emotion%20and%20cognition%20in%20fish&journal=J%20Agric%20Environ%20Ethics&volume=26&pages=7-23&publication_year=2013&author=Braithwaite%2CVA&author=Huntingford%2CF&author=Bos%2CR"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR13">Browman HI, Skiftesvik AB (2011) Welfare in aquatic organisms—is there some faith-based HAR-King going on here? Dis Aquat Org 94:255–257</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 13" href="http://scholar.google.com/scholar_lookup?&title=Welfare%20in%20aquatic%20organisms%E2%80%94is%20there%20some%20faith-based%20HAR-King%20going%20on%20here%3F&journal=Dis%20Aquat%20Org&volume=94&pages=255-257&publication_year=2011&author=Browman%2CHI&author=Skiftesvik%2CAB"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR14">Butler AB (2000) Chordate evolution and the origin of the craniates: an old brain in a new head. Anat Rec 261:111–125</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 14" href="http://scholar.google.com/scholar_lookup?&title=Chordate%20evolution%20and%20the%20origin%20of%20the%20craniates%3A%20an%20old%20brain%20in%20a%20new%20head&journal=Anat%20Rec&volume=261&pages=111-125&publication_year=2000&author=Butler%2CAB"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR15">Buxhoeveden DP, Casanova MF (2002) The minicolumn hypothesis in neuroscience. Brain 125:935–951</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 15" href="http://scholar.google.com/scholar_lookup?&title=The%20minicolumn%20hypothesis%20in%20neuroscience&journal=Brain&volume=125&pages=935-951&publication_year=2002&author=Buxhoeveden%2CDP&author=Casanova%2CMF"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR16">Cain CK, Choi J-S, LeDoux JE (2010) Active avoidance and escape learning. Encycl Behav Neurosci 1:1–9</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 16" href="http://scholar.google.com/scholar_lookup?&title=Active%20avoidance%20and%20escape%20learning&journal=Encycl%20Behav%20Neurosci&volume=1&pages=1-9&publication_year=2010&author=Cain%2CCK&author=Choi%2CJ-S&author=LeDoux%2CJE"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR17">Casanova MF (2010) Cortical organization: anatomical findings based on systems theory. Transl Neurosci 1:62–71</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 17" href="http://scholar.google.com/scholar_lookup?&title=Cortical%20organization%3A%20anatomical%20findings%20based%20on%20systems%20theory&journal=Transl%20Neurosci&volume=1&pages=62-71&publication_year=2010&author=Casanova%2CMF"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR18">Chandroo KP, Duncan IJH, Moccia RD (2004) Can fish suffer? Perspectives on sentience, pain, fear and stress. Appl Anim Behav Sci 86:225–250</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 18" href="http://scholar.google.com/scholar_lookup?&title=Can%20fish%20suffer%3F%20Perspectives%20on%20sentience%2C%20pain%2C%20fear%20and%20stress&journal=Appl%20Anim%20Behav%20Sci&volume=86&pages=225-250&publication_year=2004&author=Chandroo%2CKP&author=Duncan%2CIJH&author=Moccia%2CRD"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR19">Chen LM, Friedman RM, Ramsden BM, LaMotte RH, Roe AW (2001) Fine-scale organization of SI (area 3b) in the squirrel monkey revealed with intrinsic optical imaging. J Neurophysiol 86:3011–3029</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 19" href="http://scholar.google.com/scholar_lookup?&title=Fine-scale%20organization%20of%20SI%20%28area%203b%29%20in%20the%20squirrel%20monkey%20revealed%20with%20intrinsic%20optical%20imaging&journal=J%20Neurophysiol&volume=86&pages=3011-3029&publication_year=2001&author=Chen%2CLM&author=Friedman%2CRM&author=Ramsden%2CBM&author=LaMotte%2CRH&author=Roe%2CAW"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR20">Chen LM, Dillenburger BC, Wang F, Friedman RM, Avison MJ (2011) High-resolution functional magnetic resonance imaging of noxious heat and activations along the central sulcus in New World monkeys. Pain 152:522–532</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 20" href="http://scholar.google.com/scholar_lookup?&title=High-resolution%20functional%20magnetic%20resonance%20imaging%20of%20noxious%20heat%20and%20activations%20along%20the%20central%20sulcus%20in%20New%20World%20monkeys&journal=Pain&volume=152&pages=522-532&publication_year=2011&author=Chen%2CLM&author=Dillenburger%2CBC&author=Wang%2CF&author=Friedman%2CRM&author=Avison%2CMJ"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR21">Chervova LS (1997) Pain sensitivity of behavior of fishes. J Ichthyol 37:98–102</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 21" href="http://scholar.google.com/scholar_lookup?&title=Pain%20sensitivity%20of%20behavior%20of%20fishes&journal=J%20Ichthyol&volume=37&pages=98-102&publication_year=1997&author=Chervova%2CLS"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR22">Davis RE, Kassel J, Schwagmeyer P (1976) Telencephalic lesions and behavior in the teleost, <i>macropodus opercularis</i>: reproductive, startle reaction, and operant behavior in the male. Behav Biol 18:165–177</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 22" href="http://scholar.google.com/scholar_lookup?&title=Telencephalic%20lesions%20and%20behavior%20in%20the%20teleost%2C%20macropodus%20opercularis%3A%20reproductive%2C%20startle%20reaction%2C%20and%20operant%20behavior%20in%20the%20male&journal=Behav%20Biol&volume=18&pages=165-177&publication_year=1976&author=Davis%2CRE&author=Kassel%2CJ&author=Schwagmeyer%2CP"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR23">Demski LS (2013) The pallium and mind/behavior relationships in teleost fishes. Brain Behav Evol 82:31–44</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 23" href="http://scholar.google.com/scholar_lookup?&title=The%20pallium%20and%20mind%2Fbehavior%20relationships%20in%20teleost%20fishes&journal=Brain%20Behav%20Evol&volume=82&pages=31-44&publication_year=2013&author=Demski%2CLS"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR24">Dimitrijevic MR, Nathan PW (1968) Studies of spasticity in man. 3. Analysis of reflex activity evoked by noxious cutaneous stimulation. Brain 91:349–368</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 24" href="http://scholar.google.com/scholar_lookup?&title=Studies%20of%20spasticity%20in%20man.%203.%20Analysis%20of%20reflex%20activity%20evoked%20by%20noxious%20cutaneous%20stimulation&journal=Brain&volume=91&pages=349-368&publication_year=1968&author=Dimitrijevic%2CMR&author=Nathan%2CPW"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR25">Dugas-Ford J, Rowell JJ, Ragsdale CW (2012) Cell-type homologies and the origins of the neocortex. PNAS 109:16974–16979</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 25" href="http://scholar.google.com/scholar_lookup?&title=Cell-type%20homologies%20and%20the%20origins%20of%20the%20neocortex&journal=PNAS&volume=109&pages=16974-16979&publication_year=2012&author=Dugas-Ford%2CJ&author=Rowell%2CJJ&author=Ragsdale%2CCW"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR26">Dunlop R, Laming P (2005) Mechanoreceptive and nociceptive responses in the central nervous system of goldfish (<i>Carassius auratus</i>) and trout (<i>Oncorrhynchus mykiss</i>). J Pain 6:561–568</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 26" href="http://scholar.google.com/scholar_lookup?&title=Mechanoreceptive%20and%20nociceptive%20responses%20in%20the%20central%20nervous%20system%20of%20goldfish%20%28Carassius%20auratus%29%20and%20trout%20%28Oncorrhynchus%20mykiss%29&journal=J%20Pain&volume=6&pages=561-568&publication_year=2005&author=Dunlop%2CR&author=Laming%2CP"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR27">Dunlop R, Millsopp S, Laming P (2006) Avoidance learning in goldfish (<i>Carassius auratus</i>) and implications for pain perception. Appl Anim Behav Sci 976:255–271</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 27" href="http://scholar.google.com/scholar_lookup?&title=Avoidance%20learning%20in%20goldfish%20%28Carassius%20auratus%29%20and%20implications%20for%20pain%20perception&journal=Appl%20Anim%20Behav%20Sci&volume=976&pages=255-271&publication_year=2006&author=Dunlop%2CR&author=Millsopp%2CS&author=Laming%2CP"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR28">Echleter SM, Saidel WM (1981) Forebrain connections in the goldfish support telencephalic homologies with land vertebrates. Science 212:683–685</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 28" href="http://scholar.google.com/scholar_lookup?&title=Forebrain%20connections%20in%20the%20goldfish%20support%20telencephalic%20homologies%20with%20land%20vertebrates&journal=Science&volume=212&pages=683-685&publication_year=1981&author=Echleter%2CSM&author=Saidel%2CWM"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR29">Ellis T, Oidtmann B, St Hilaire S, Turnbull JF, North BP, MacIntyre CM, Nikolaidis J, Hoyle I, Kestin SC, Knowles TG (2008) Fin erosion in farmed fish. In: Branson EJ (ed) Fish welfare, chapter 9. Wiley, New York. ISBN 978-0-470-69804-4</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 29" href="http://scholar.google.com/scholar_lookup?&title=Fin%20erosion%20in%20farmed%20fish&publication_year=2008&author=Ellis%2CT&author=Oidtmann%2CB&author=St%20Hilaire%2CS&author=Turnbull%2CJF&author=North%2CBP&author=MacIntyre%2CCM&author=Nikolaidis%2CJ&author=Hoyle%2CI&author=Kestin%2CSC&author=Knowles%2CTG"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR30">Fatira E, Papandroulakis N, Pavlidis M (2014) Diel changes in plasma cortisol and effects of size and stress duration on the cortisol response in European sea bass (<i>Dicentrarchus labrax</i>). Fish Physiol Biochem. doi:<a href="https://doi.org/10.1007/s10695-013-9896-1" data-track="click_references" data-track-action="external reference" data-track-value="external reference" data-track-label="10.1007/s10695-013-9896-1">10.1007/s10695-013-9896-1</a> </p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 30" href="http://scholar.google.com/scholar_lookup?&title=Diel%20changes%20in%20plasma%20cortisol%20and%20effects%20of%20size%20and%20stress%20duration%20on%20the%20cortisol%20response%20in%20European%20sea%20bass%20%28Dicentrarchus%20labrax%29&journal=Fish%20Physiol%20Biochem&doi=10.1007%2Fs10695-013-9896-1&publication_year=2014&author=Fatira%2CE&author=Papandroulakis%2CN&author=Pavlidis%2CM"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR31">Filk G, Klaren PHM, Van den Burg EH, Metz JR, Huising MO (2006) CRF and stress in fish. Gen Comp Endocrinol 146:36–44</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 31" href="http://scholar.google.com/scholar_lookup?&title=CRF%20and%20stress%20in%20fish&journal=Gen%20Comp%20Endocrinol&volume=146&pages=36-44&publication_year=2006&author=Filk%2CG&author=Klaren%2CPHM&author=Burg%2CEH&author=Metz%2CJR&author=Huising%2CMO"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR32">Flood NC, Overmier JB, Savage GE (1976) Teleost telencephalon and learning: an interpretive review of data and hypotheses. Physiol Behav 16:783–798</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 32" href="http://scholar.google.com/scholar_lookup?&title=Teleost%20telencephalon%20and%20learning%3A%20an%20interpretive%20review%20of%20data%20and%20hypotheses&journal=Physiol%20Behav&volume=16&pages=783-798&publication_year=1976&author=Flood%2CNC&author=Overmier%2CJB&author=Savage%2CGE"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR33">Fu C, Cao ZD, Fu SJ (2013) The effects of caudal fin amputation on metabolic interaction between digestion and locomotion in juveniles of three cyprinid fish species with different metabolic modes. Comp Biochem Physiol A: Mol Integr Physiol 164:456–465</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 33" href="http://scholar.google.com/scholar_lookup?&title=The%20effects%20of%20caudal%20fin%20amputation%20on%20metabolic%20interaction%20between%20digestion%20and%20locomotion%20in%20juveniles%20of%20three%20cyprinid%20fish%20species%20with%20different%20metabolic%20modes&journal=Comp%20Biochem%20Physiol%20A%3A%20Mol%20Integr%20Physiol&volume=164&pages=456-465&publication_year=2013&author=Fu%2CC&author=Cao%2CZD&author=Fu%2CSJ"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR34">Giassi ACC, Ellis W, Maler L (2012) Organization of the gymnotiform fish pallium in relation to learning and memory: III. Intrinsic connections. J Comp Neurol 520:3369–3394</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 34" href="http://scholar.google.com/scholar_lookup?&title=Organization%20of%20the%20gymnotiform%20fish%20pallium%20in%20relation%20to%20learning%20and%20memory%3A%20III.%20Intrinsic%20connections&journal=J%20Comp%20Neurol&volume=520&pages=3369-3394&publication_year=2012&author=Giassi%2CACC&author=Ellis%2CW&author=Maler%2CL"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR35">Guerrini R, Parrini E (2010) Neuronal migration disorders. Neurobiol Dis 38:154–166</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 35" href="http://scholar.google.com/scholar_lookup?&title=Neuronal%20migration%20disorders&journal=Neurobiol%20Dis&volume=38&pages=154-166&publication_year=2010&author=Guerrini%2CR&author=Parrini%2CE"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR36">Guerrini R, Dobyns WB, Barkovich AJ (2008) Abnormal development of the human cerebral cortex: genetics, functional consequences and treatment options. TINS 31:154–162</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 36" href="http://scholar.google.com/scholar_lookup?&title=Abnormal%20development%20of%20the%20human%20cerebral%20cortex%3A%20genetics%2C%20functional%20consequences%20and%20treatment%20options&journal=TINS&volume=31&pages=154-162&publication_year=2008&author=Guerrini%2CR&author=Dobyns%2CWB&author=Barkovich%2CAJ"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR37">Gunturkun O (2005) The avian ‘prefrontal cortex’ and cognition. Curr Opin Neurobiol 15:686–693</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 37" href="http://scholar.google.com/scholar_lookup?&title=The%20avian%20%E2%80%98prefrontal%20cortex%E2%80%99%20and%20cognition&journal=Curr%20Opin%20Neurobiol&volume=15&pages=686-693&publication_year=2005&author=Gunturkun%2CO"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR38">Gunturkun O (2012) The convergent evolution of neural substrates for cognition. Psychol Res 76:212–219</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 38" href="http://scholar.google.com/scholar_lookup?&title=The%20convergent%20evolution%20of%20neural%20substrates%20for%20cognition&journal=Psychol%20Res&volume=76&pages=212-219&publication_year=2012&author=Gunturkun%2CO"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR39">Hainsworth FR, Overmier JB, Snowdon CT (1967) Specific and permanent deficits in instrumental avoidance responding following forebrain ablation in the goldfish. J Comp Physiol Psychol 63:111–116</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 39" href="http://scholar.google.com/scholar_lookup?&title=Specific%20and%20permanent%20deficits%20in%20instrumental%20avoidance%20responding%20following%20forebrain%20ablation%20in%20the%20goldfish&journal=J%20Comp%20Physiol%20Psychol&volume=63&pages=111-116&publication_year=1967&author=Hainsworth%2CFR&author=Overmier%2CJB&author=Snowdon%2CCT"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR40">Hjorth JT, Key B (2001) Are pioneer axons guided by regulatory gene expression domains in the zebrafish forebrain? High-resolution analysis of the patterning of the zebrafish brain during axon tract formation. Dev Biol 229:271–286</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 40" href="http://scholar.google.com/scholar_lookup?&title=Are%20pioneer%20axons%20guided%20by%20regulatory%20gene%20expression%20domains%20in%20the%20zebrafish%20forebrain%3F%20High-resolution%20analysis%20of%20the%20patterning%20of%20the%20zebrafish%20brain%20during%20axon%20tract%20formation&journal=Dev%20Biol&volume=229&pages=271-286&publication_year=2001&author=Hjorth%2CJT&author=Key%2CB"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR41">Hurtado-Parrado C (2010) Neuronal mechanisms of learning in teleost fish. Univ Psychol 9:663–672</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 41" href="http://scholar.google.com/scholar_lookup?&title=Neuronal%20mechanisms%20of%20learning%20in%20teleost%20fish&journal=Univ%20Psychol&volume=9&pages=663-672&publication_year=2010&author=Hurtado-Parrado%2CC"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR42">Ingle D (1973) Evolutionary perspectives on the function of the optic tectum. Brain Behav Evol 8:211–237</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 42" href="http://scholar.google.com/scholar_lookup?&title=Evolutionary%20perspectives%20on%20the%20function%20of%20the%20optic%20tectum&journal=Brain%20Behav%20Evol&volume=8&pages=211-237&publication_year=1973&author=Ingle%2CD"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR43">Jacobs C, de Gaff TA, Goebel R, Sack AT (2012) The temporal dynamics of early visual cortex involvement in behavioral priming. PLoS ONE 7:e48808</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 43" href="http://scholar.google.com/scholar_lookup?&title=The%20temporal%20dynamics%20of%20early%20visual%20cortex%20involvement%20in%20behavioral%20priming&journal=PLoS%20ONE&volume=7&publication_year=2012&author=Jacobs%2CC&author=Gaff%2CTA&author=Goebel%2CR&author=Sack%2CAT"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR44">Jarvis ED, Yu J, Rivas MV, Horita H, Feenders G, Whitney O, Jarvis SC, Jarvis ER, Kubikova L, Puck AEP, Siang-Bakshi C, Martin S, McElroy M, Hara E, Howard J, Pfenning A, Mouritsen H, Chen C-C, Wada K (2013) Global view of the functional molecular organization of the avian cerebrum: mirror images and functional columns. J Comp Neurol 521:3614–3665</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 44" href="http://scholar.google.com/scholar_lookup?&title=Global%20view%20of%20the%20functional%20molecular%20organization%20of%20the%20avian%20cerebrum%3A%20mirror%20images%20and%20functional%20columns&journal=J%20Comp%20Neurol&volume=521&pages=3614-3665&publication_year=2013&author=Jarvis%2CED&author=Yu%2CJ&author=Rivas%2CMV&author=Horita%2CH&author=Feenders%2CG&author=Whitney%2CO&author=Jarvis%2CSC&author=Jarvis%2CER&author=Kubikova%2CL&author=Puck%2CAEP&author=Siang-Bakshi%2CC&author=Martin%2CS&author=McElroy%2CM&author=Hara%2CE&author=Howard%2CJ&author=Pfenning%2CA&author=Mouritsen%2CH&author=Chen%2CC-C&author=Wada%2CK"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR45">Jones RC (2013) Science, sentience, and animal welfare. Biol Philos 28:1–30</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 45" href="http://scholar.google.com/scholar_lookup?&title=Science%2C%20sentience%2C%20and%20animal%20welfare&journal=Biol%20Philos&volume=28&pages=1-30&publication_year=2013&author=Jones%2CRC"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR46">Jones AKP, Friston KJ, Qi LY, Harris M, Cunningham VJ, Jones T, Feinman C, Frackowiak RSJ (1991) Sites of action of morphine in the brain. Lancet 338:825</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 46" href="http://scholar.google.com/scholar_lookup?&title=Sites%20of%20action%20of%20morphine%20in%20the%20brain&journal=Lancet&volume=338&publication_year=1991&author=Jones%2CAKP&author=Friston%2CKJ&author=Qi%2CLY&author=Harris%2CM&author=Cunningham%2CVJ&author=Jones%2CT&author=Feinman%2CC&author=Frackowiak%2CRSJ"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR47">Jones SG, Kamunde C, Lemke K, Stevens ED (2012) The dose-response relation for the antinociceptive effect of morphine in a fish, rainbow trout. J Vet Pharmacol Ther 35:563–570</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 47" href="http://scholar.google.com/scholar_lookup?&title=The%20dose-response%20relation%20for%20the%20antinociceptive%20effect%20of%20morphine%20in%20a%20fish%2C%20rainbow%20trout&journal=J%20Vet%20Pharmacol%20Ther&volume=35&pages=563-570&publication_year=2012&author=Jones%2CSG&author=Kamunde%2CC&author=Lemke%2CK&author=Stevens%2CED"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR48">Kaas JH (1997) Topographic maps are fundamental to sensory processing. Brain Res Bull 44:107–112</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 48" href="http://scholar.google.com/scholar_lookup?&title=Topographic%20maps%20are%20fundamental%20to%20sensory%20processing&journal=Brain%20Res%20Bull&volume=44&pages=107-112&publication_year=1997&author=Kaas%2CJH"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR49">Kaas JH (2012) Evolution of columns, modules, and domains in the neocortex of primates. PNAS 109:10655–10660</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 49" href="http://scholar.google.com/scholar_lookup?&title=Evolution%20of%20columns%2C%20modules%2C%20and%20domains%20in%20the%20neocortex%20of%20primates&journal=PNAS&volume=109&pages=10655-10660&publication_year=2012&author=Kaas%2CJH"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR50">Kanai R, Tsuchiya N (2012) Qualia. Curr Biol 22:R392–R396</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 50" href="http://scholar.google.com/scholar_lookup?&title=Qualia&journal=Curr%20Biol&volume=22&pages=R392-R396&publication_year=2012&author=Kanai%2CR&author=Tsuchiya%2CN"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR51">Kaplan H, Aronson LR (1967) Effect of forebrain ablation on the performance of a conditioned avoidance response in the teleost fish, <i>Tilapia H.</i> Macrocephala. Anim Behav 15:438–448</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 51" href="http://scholar.google.com/scholar_lookup?&title=Effect%20of%20forebrain%20ablation%20on%20the%20performance%20of%20a%20conditioned%20avoidance%20response%20in%20the%20teleost%20fish%2C%20Tilapia%20H.%20Macrocephala&journal=Anim%20Behav&volume=15&pages=438-448&publication_year=1967&author=Kaplan%2CH&author=Aronson%2CLR"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR52">Kavaliers M (1988) Evolutionary and comparative aspects of nociception. Brain Res Bull 21:923–931</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 52" href="http://scholar.google.com/scholar_lookup?&title=Evolutionary%20and%20comparative%20aspects%20of%20nociception&journal=Brain%20Res%20Bull&volume=21&pages=923-931&publication_year=1988&author=Kavaliers%2CM"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR53">Kirsch JA, Gunturkun O, Rose J (2008) Insight without cortex: lessons from the avian brain. Conscious Cogn 17:475–483</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 53" href="http://scholar.google.com/scholar_lookup?&title=Insight%20without%20cortex%3A%20lessons%20from%20the%20avian%20brain&journal=Conscious%20Cogn&volume=17&pages=475-483&publication_year=2008&author=Kirsch%2CJA&author=Gunturkun%2CO&author=Rose%2CJ"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR130">Kittilsen S (2013) Functional aspects of emotions in fish. Behav Process 100:153–159</p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR54">Koivisto M, Silvanto J (2012) Visual feature binding: the critical time windows of V1/V2 and parietal activity. Neuroimage 59:1608–1614</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 55" href="http://scholar.google.com/scholar_lookup?&title=Visual%20feature%20binding%3A%20the%20critical%20time%20windows%20of%20V1%2FV2%20and%20parietal%20activity&journal=Neuroimage&volume=59&pages=1608-1614&publication_year=2012&author=Koivisto%2CM&author=Silvanto%2CJ"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR55">Koivisto M, Mantyla T, Silvanto J (2010) The role of early visual corex (V1/V2) in conscious and unconscious visual perception. Neuroimage 51:828–834</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 56" href="http://scholar.google.com/scholar_lookup?&title=The%20role%20of%20early%20visual%20corex%20%28V1%2FV2%29%20in%20conscious%20and%20unconscious%20visual%20perception&journal=Neuroimage&volume=51&pages=828-834&publication_year=2010&author=Koivisto%2CM&author=Mantyla%2CT&author=Silvanto%2CJ"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR56">Koivisto M, Railo H, Salminen-Vaparanta N (2011) Transcranial magnetic stimulation of early visual cortex interferes with subjective visual awareness and objective forced-choice performance. Conscious Cogn 20:288–298</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 57" href="http://scholar.google.com/scholar_lookup?&title=Transcranial%20magnetic%20stimulation%20of%20early%20visual%20cortex%20interferes%20with%20subjective%20visual%20awareness%20and%20objective%20forced-choice%20performance&journal=Conscious%20Cogn&volume=20&pages=288-298&publication_year=2011&author=Koivisto%2CM&author=Railo%2CH&author=Salminen-Vaparanta%2CN"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR57">Krahe R, Maler L (2014) Neural maps in the electrosensory system of weakly electric fish. Curr Opin Neurobiol 24:13–21</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 58" href="http://scholar.google.com/scholar_lookup?&title=Neural%20maps%20in%20the%20electrosensory%20system%20of%20weakly%20electric%20fish&journal=Curr%20Opin%20Neurobiol&volume=24&pages=13-21&publication_year=2014&author=Krahe%2CR&author=Maler%2CL"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR58">Kupers R, Pitto M (2013) Compensatory plasticity and cross-modal reorganization following early visual deprivation. Neurosci Biobehav Rev. doi:<a href="https://doi.org/10.1016/j.neubiorev.2013.08.001" data-track="click_references" data-track-action="external reference" data-track-value="external reference" data-track-label="10.1016/j.neubiorev.2013.08.001">10.1016/j.neubiorev.2013.08.001</a> </p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 59" href="http://scholar.google.com/scholar_lookup?&title=Compensatory%20plasticity%20and%20cross-modal%20reorganization%20following%20early%20visual%20deprivation&journal=Neurosci%20Biobehav%20Rev&doi=10.1016%2Fj.neubiorev.2013.08.001&publication_year=2013&author=Kupers%2CR&author=Pitto%2CM"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR59">Larkum M (2013) A cellular mechanism for cortical associations: an organizing principle for the cerebral cortex. TINS 36:141–151</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 60" href="http://scholar.google.com/scholar_lookup?&title=A%20cellular%20mechanism%20for%20cortical%20associations%3A%20an%20organizing%20principle%20for%20the%20cerebral%20cortex&journal=TINS&volume=36&pages=141-151&publication_year=2013&author=Larkum%2CM"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR60">Luque MA, Perez-Perez MP, Herrero L, Torres B (2005) Involvement of the optic tectum and mesencephalic reticular formation in the generation of saccadic eye movements in goldfish. Brain Res Rev 49:388–397</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 61" href="http://scholar.google.com/scholar_lookup?&title=Involvement%20of%20the%20optic%20tectum%20and%20mesencephalic%20reticular%20formation%20in%20the%20generation%20of%20saccadic%20eye%20movements%20in%20goldfish&journal=Brain%20Res%20Rev&volume=49&pages=388-397&publication_year=2005&author=Luque%2CMA&author=Perez-Perez%2CMP&author=Herrero%2CL&author=Torres%2CB"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR61">Maier A, Adams GK, Aura C, Leopold DA (2010) Distinct superficial and deep laminar domains of activity in the visual cortex during rest and stimulation. Front Syst Neurosci. doi:<a href="https://doi.org/10.3389/fnsys.2010.00031" data-track="click_references" data-track-action="external reference" data-track-value="external reference" data-track-label="10.3389/fnsys.2010.00031">10.3389/fnsys.2010.00031</a> </p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 62" href="http://scholar.google.com/scholar_lookup?&title=Distinct%20superficial%20and%20deep%20laminar%20domains%20of%20activity%20in%20the%20visual%20cortex%20during%20rest%20and%20stimulation&journal=Front%20Syst%20Neurosci&doi=10.3389%2Ffnsys.2010.00031&publication_year=2010&author=Maier%2CA&author=Adams%2CGK&author=Aura%2CC&author=Leopold%2CDA"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR62">Malafoglia V, Bryant B, Raffaeli W, Giordano A, Bellipanni G (2013) The zebrafish as a model for nociception studies. J Cell Physiol 228:1956–1966</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 63" href="http://scholar.google.com/scholar_lookup?&title=The%20zebrafish%20as%20a%20model%20for%20nociception%20studies&journal=J%20Cell%20Physiol&volume=228&pages=1956-1966&publication_year=2013&author=Malafoglia%2CV&author=Bryant%2CB&author=Raffaeli%2CW&author=Giordano%2CA&author=Bellipanni%2CG"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR63">Mancini F, Haggard P, Iannetti GD, Longo MR, Sereno MI (2012) Fine-grained nociceptive maps in primary somatosensory cortex. J Neurosci 32:17155–17162</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 64" href="http://scholar.google.com/scholar_lookup?&title=Fine-grained%20nociceptive%20maps%20in%20primary%20somatosensory%20cortex&journal=J%20Neurosci&volume=32&pages=17155-17162&publication_year=2012&author=Mancini%2CF&author=Haggard%2CP&author=Iannetti%2CGD&author=Longo%2CMR&author=Sereno%2CMI"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR64">Manek AK, Ferrari MC, Niyogi S, Chivers DP (2014) The interactive effects of multiple stressors on physiological stress responders and club cell investment in fathead minnows. Sci Total Environ 467–477:90–97</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 65" href="http://scholar.google.com/scholar_lookup?&title=The%20interactive%20effects%20of%20multiple%20stressors%20on%20physiological%20stress%20responders%20and%20club%20cell%20investment%20in%20fathead%20minnows&journal=Sci%20Total%20Environ&volume=467%E2%80%93477&pages=90-97&publication_year=2014&author=Manek%2CAK&author=Ferrari%2CMC&author=Niyogi%2CS&author=Chivers%2CDP"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR65">Manger PR, Elston GN, Pettigrew JD (2002) Multiple maps and activity-dependent representational plasticity in the anterior Wust of the adult barn owl (<i>Tyto alba</i>). Eur J Neurosci 16:743–750</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 66" href="http://scholar.google.com/scholar_lookup?&title=Multiple%20maps%20and%20activity-dependent%20representational%20plasticity%20in%20the%20anterior%20Wust%20of%20the%20adult%20barn%20owl%20%28Tyto%20alba%29&journal=Eur%20J%20Neurosci&volume=16&pages=743-750&publication_year=2002&author=Manger%2CPR&author=Elston%2CGN&author=Pettigrew%2CJD"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR66">Matthies BK, Franklin KBJ (1992) Formalin pain is expressed in decerebrate rats but not attenuated by morphine. Pain 51:199–206</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 67" href="http://scholar.google.com/scholar_lookup?&title=Formalin%20pain%20is%20expressed%20in%20decerebrate%20rats%20but%20not%20attenuated%20by%20morphine&journal=Pain&volume=51&pages=199-206&publication_year=1992&author=Matthies%2CBK&author=Franklin%2CKBJ"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR67">Matthies BK, Franklin KBJ (1995) Effects of partial decortication on the opioid analgesia in the formalin test. Behav Brain Res 67:59–66</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 68" href="http://scholar.google.com/scholar_lookup?&title=Effects%20of%20partial%20decortication%20on%20the%20opioid%20analgesia%20in%20the%20formalin%20test&journal=Behav%20Brain%20Res&volume=67&pages=59-66&publication_year=1995&author=Matthies%2CBK&author=Franklin%2CKBJ"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR68">Mazzola L, Faillenot I, Barral F-G, Mauguiere F, Peyron R (2012) Spatial segregation of somatosensory and pain activations in the human operculo-insular cortex. Neuroimage 60:5409–5418</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 69" href="http://scholar.google.com/scholar_lookup?&title=Spatial%20segregation%20of%20somatosensory%20and%20pain%20activations%20in%20the%20human%20operculo-insular%20cortex&journal=Neuroimage&volume=60&pages=5409-5418&publication_year=2012&author=Mazzola%2CL&author=Faillenot%2CI&author=Barral%2CF-G&author=Mauguiere%2CF&author=Peyron%2CR"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR69">Meek J (1983) Functional anatomy of the tectum mesencephala of the goldfish. An explorative analysis of the functional implications of the laminar structural organization of the tectum. Brain Res 287:247–297</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 70" href="http://scholar.google.com/scholar_lookup?&title=Functional%20anatomy%20of%20the%20tectum%20mesencephala%20of%20the%20goldfish.%20An%20explorative%20analysis%20of%20the%20functional%20implications%20of%20the%20laminar%20structural%20organization%20of%20the%20tectum&journal=Brain%20Res&volume=287&pages=247-297&publication_year=1983&author=Meek%2CJ"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR70">Mountcastle VB (1997) The columnar organization of the neocortex. Brain 120:701–722</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 71" href="http://scholar.google.com/scholar_lookup?&title=The%20columnar%20organization%20of%20the%20neocortex&journal=Brain&volume=120&pages=701-722&publication_year=1997&author=Mountcastle%2CVB"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR71">Mueller T, Wullimann MF (2009) An evolutionary interpretation of teleostean forebrain anatomy. Brain Behav Evol 74:30–42</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 72" href="http://scholar.google.com/scholar_lookup?&title=An%20evolutionary%20interpretation%20of%20teleostean%20forebrain%20anatomy&journal=Brain%20Behav%20Evol&volume=74&pages=30-42&publication_year=2009&author=Mueller%2CT&author=Wullimann%2CMF"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR72">Mueller T, Dong Z, Berberoglu MA, Guo S (2011) The dorsal pallium in zebrafish, <i>Danio rerio</i> (Cyprinidae, Teleostei). Brain Res 1381:95–105</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 73" href="http://scholar.google.com/scholar_lookup?&title=The%20dorsal%20pallium%20in%20zebrafish%2C%20Danio%20rerio%20%28Cyprinidae%2C%20Teleostei%29&journal=Brain%20Res&volume=1381&pages=95-105&publication_year=2011&author=Mueller%2CT&author=Dong%2CZ&author=Berberoglu%2CMA&author=Guo%2CS"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR73">Nguyen AP, Spetch ML, Crowder NA, Winship IR, Hurd PL, Wylie DR (2004) A dissociation of motion and spatial-pattern vision in the avian telencephalon: implications for the evolution of “visual streams”. J Neurosci 24:4962–4970</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 74" href="http://scholar.google.com/scholar_lookup?&title=A%20dissociation%20of%20motion%20and%20spatial-pattern%20vision%20in%20the%20avian%20telencephalon%3A%20implications%20for%20the%20evolution%20of%20%E2%80%9Cvisual%20streams%E2%80%9D&journal=J%20Neurosci&volume=24&pages=4962-4970&publication_year=2004&author=Nguyen%2CAP&author=Spetch%2CML&author=Crowder%2CNA&author=Winship%2CIR&author=Hurd%2CPL&author=Wylie%2CDR"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR74">Nieuwenhuys R (1994) The neocortex. An overview of its evolutionary development, structural organization and synaptology. Anat Embryol 190:307–337</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 75" href="http://scholar.google.com/scholar_lookup?&title=The%20neocortex.%20An%20overview%20of%20its%20evolutionary%20development%2C%20structural%20organization%20and%20synaptology&journal=Anat%20Embryol&volume=190&pages=307-337&publication_year=1994&author=Nieuwenhuys%2CR"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR75">Nordgreen J, Horsberg TE, Ranheim B, Chen ACN (2007) Somatosensory evoked potentials in the telencephalon of Atlanic salmon (<i>Salmo salar</i>) following galvanic stimulation of the tail. J Comp Physiol A 193:1235–1242</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 76" href="http://scholar.google.com/scholar_lookup?&title=Somatosensory%20evoked%20potentials%20in%20the%20telencephalon%20of%20Atlanic%20salmon%20%28Salmo%20salar%29%20following%20galvanic%20stimulation%20of%20the%20tail&journal=J%20Comp%20Physiol%20A&volume=193&pages=1235-1242&publication_year=2007&author=Nordgreen%2CJ&author=Horsberg%2CTE&author=Ranheim%2CB&author=Chen%2CACN"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR76">Northcutt RG (2008) Forebrain evolution in bony fishes. Brain Res Bull 75:191–205</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 77" href="http://scholar.google.com/scholar_lookup?&title=Forebrain%20evolution%20in%20bony%20fishes&journal=Brain%20Res%20Bull&volume=75&pages=191-205&publication_year=2008&author=Northcutt%2CRG"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR77">Northcutt RG (2011) Do teleost fishes possess a homolog of mammalian isocortex? Brain Behav Evol 78:136–138</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 78" href="http://scholar.google.com/scholar_lookup?&title=Do%20teleost%20fishes%20possess%20a%20homolog%20of%20mammalian%20isocortex%3F&journal=Brain%20Behav%20Evol&volume=78&pages=136-138&publication_year=2011&author=Northcutt%2CRG"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR78">Northcutt RG, Plassman W, Holmes PH, Saidel WM (2004) A pallial visual area in the telencephalon of the bony fish <i>Polypterus</i>. Brain Behav Evol 64:1–10</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 79" href="http://scholar.google.com/scholar_lookup?&title=A%20pallial%20visual%20area%20in%20the%20telencephalon%20of%20the%20bony%20fish%20Polypterus&journal=Brain%20Behav%20Evol&volume=64&pages=1-10&publication_year=2004&author=Northcutt%2CRG&author=Plassman%2CW&author=Holmes%2CPH&author=Saidel%2CWM"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR79">Ohiorhenuan IE, Mechler F, Purpura KP, Schmid AM, Hu Q, Victor JD (2010) Sparse coding and high-order correlations in fine-scale cortical networks. Nature 466:617–622</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 80" href="http://scholar.google.com/scholar_lookup?&title=Sparse%20coding%20and%20high-order%20correlations%20in%20fine-scale%20cortical%20networks&journal=Nature&volume=466&pages=617-622&publication_year=2010&author=Ohiorhenuan%2CIE&author=Mechler%2CF&author=Purpura%2CKP&author=Schmid%2CAM&author=Hu%2CQ&author=Victor%2CJD"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR80">Opris I (2013) Inter-laminar microcircuits across neocortex: repair and augmentation. Front Syst Neurosci 7:80</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 81" href="http://scholar.google.com/scholar_lookup?&title=Inter-laminar%20microcircuits%20across%20neocortex%3A%20repair%20and%20augmentation&journal=Front%20Syst%20Neurosci&volume=7&publication_year=2013&author=Opris%2CI"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR81">Opris I, Fuqua JL, Huetti PF, Gerhardt GA, Berger TW, Hampson RE, Deadwyler SA (2012) Closing the loop in primate prefrontal cortex: inter-laminar processing. Front Neural Circuits 6:1–13</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 82" href="http://scholar.google.com/scholar_lookup?&title=Closing%20the%20loop%20in%20primate%20prefrontal%20cortex%3A%20inter-laminar%20processing&journal=Front%20Neural%20Circuits&volume=6&pages=1-13&publication_year=2012&author=Opris%2CI&author=Fuqua%2CJL&author=Huetti%2CPF&author=Gerhardt%2CGA&author=Berger%2CTW&author=Hampson%2CRE&author=Deadwyler%2CSA"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR82">Overmier JB, Gross D (1974) Effects of telencephalic ablation upon nest-building and avoidance bahaviors in East African mouthbreeding fish, <i>Tilapia mossambica</i>. Behav Biol 12:211–222</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 83" href="http://scholar.google.com/scholar_lookup?&title=Effects%20of%20telencephalic%20ablation%20upon%20nest-building%20and%20avoidance%20bahaviors%20in%20East%20African%20mouthbreeding%20fish%2C%20Tilapia%20mossambica&journal=Behav%20Biol&volume=12&pages=211-222&publication_year=1974&author=Overmier%2CJB&author=Gross%2CD"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR83">Overmier JB, Papini MR (1985) Serial ablations of the telencephalon and avoidance learning by goldfish (<i>Carassius auratus</i>). Behav Neurosci 99:509–520</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 84" href="http://scholar.google.com/scholar_lookup?&title=Serial%20ablations%20of%20the%20telencephalon%20and%20avoidance%20learning%20by%20goldfish%20%28Carassius%20auratus%29&journal=Behav%20Neurosci&volume=99&pages=509-520&publication_year=1985&author=Overmier%2CJB&author=Papini%2CMR"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR84">Perez-Perez MP, Luque MA, Herrero L, Nunez-Abades PA, Torres B (2003) Connectivity of the goldfish optic tectum with the mesencephalic and rhombencephalic reticular formation. Exp Brain Res 151:123–135</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 85" href="http://scholar.google.com/scholar_lookup?&title=Connectivity%20of%20the%20goldfish%20optic%20tectum%20with%20the%20mesencephalic%20and%20rhombencephalic%20reticular%20formation&journal=Exp%20Brain%20Res&volume=151&pages=123-135&publication_year=2003&author=Perez-Perez%2CMP&author=Luque%2CMA&author=Herrero%2CL&author=Nunez-Abades%2CPA&author=Torres%2CB"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR85">Pert A, Yaksh T (1975) Localization of the antinociceptive action of morphine in primate brain. Pharmacol Biochem Behav 3:133–138</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 86" href="http://scholar.google.com/scholar_lookup?&title=Localization%20of%20the%20antinociceptive%20action%20of%20morphine%20in%20primate%20brain&journal=Pharmacol%20Biochem%20Behav&volume=3&pages=133-138&publication_year=1975&author=Pert%2CA&author=Yaksh%2CT"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR86">Pietri T, Manalo E, Ryan J, Saint-Amant L, Washbourne P (2009) Glutamate drives the touch response through a rostral loop in the spinal cord of zebrafish embryos. Dev Neurobiol 69:780–795</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 87" href="http://scholar.google.com/scholar_lookup?&title=Glutamate%20drives%20the%20touch%20response%20through%20a%20rostral%20loop%20in%20the%20spinal%20cord%20of%20zebrafish%20embryos&journal=Dev%20Neurobiol&volume=69&pages=780-795&publication_year=2009&author=Pietri%2CT&author=Manalo%2CE&author=Ryan%2CJ&author=Saint-Amant%2CL&author=Washbourne%2CP"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR87">Pollen DA (2011) On the emergence of primary visual perception. Cereb Cortex 21:1941–1953</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 88" href="http://scholar.google.com/scholar_lookup?&title=On%20the%20emergence%20of%20primary%20visual%20perception&journal=Cereb%20Cortex&volume=21&pages=1941-1953&publication_year=2011&author=Pollen%2CDA"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR88">Portavella M, Vargas JP (2005) Emotional and spatial learning in goldfish is dependent on different telencephalic pallial systems. Eur J Neurosci 21:2800–2806</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 89" href="http://scholar.google.com/scholar_lookup?&title=Emotional%20and%20spatial%20learning%20in%20goldfish%20is%20dependent%20on%20different%20telencephalic%20pallial%20systems&journal=Eur%20J%20Neurosci&volume=21&pages=2800-2806&publication_year=2005&author=Portavella%2CM&author=Vargas%2CJP"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR89">Portavella M, Salas CM, Vargas JP, Papini MR (2003) Involvement of the telencephalon in spaced-trial avoidance learning in the goldfish (<i>Carassius auratus</i>). Physiol Behav 80:49–56</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 90" href="http://scholar.google.com/scholar_lookup?&title=Involvement%20of%20the%20telencephalon%20in%20spaced-trial%20avoidance%20learning%20in%20the%20goldfish%20%28Carassius%20auratus%29&journal=Physiol%20Behav&volume=80&pages=49-56&publication_year=2003&author=Portavella%2CM&author=Salas%2CCM&author=Vargas%2CJP&author=Papini%2CMR"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR90">Portavella M, Torres B, Salas C (2004a) Avoidance response in goldfish: emotional and temporal involvement of medial and lateral telencephalic pallium. J Neurosci 24:2335–2342</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 91" href="http://scholar.google.com/scholar_lookup?&title=Avoidance%20response%20in%20goldfish%3A%20emotional%20and%20temporal%20involvement%20of%20medial%20and%20lateral%20telencephalic%20pallium&journal=J%20Neurosci&volume=24&pages=2335-2342&publication_year=2004&author=Portavella%2CM&author=Torres%2CB&author=Salas%2CC"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR91">Portavella M, Torres B, Salas C, Papini MR (2004b) Lesions of the medial pallium, but not of the lateral pallium, disrupt spaced-trial avoidance learning in goldfish (<i>Carassius auratus</i>). Neurosci Lett 362:75–78</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 92" href="http://scholar.google.com/scholar_lookup?&title=Lesions%20of%20the%20medial%20pallium%2C%20but%20not%20of%20the%20lateral%20pallium%2C%20disrupt%20spaced-trial%20avoidance%20learning%20in%20goldfish%20%28Carassius%20auratus%29&journal=Neurosci%20Lett&volume=362&pages=75-78&publication_year=2004&author=Portavella%2CM&author=Torres%2CB&author=Salas%2CC&author=Papini%2CMR"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR92">Precht JC, von der Emde G, Wolfart J, Karamursel S, Akoev GN, Andrianov YN, Bullock TH (1998) Sensory processing in the pallium of a Mormyrid fish. J Neurosci 18:7381–7393</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 93" href="http://scholar.google.com/scholar_lookup?&title=Sensory%20processing%20in%20the%20pallium%20of%20a%20Mormyrid%20fish&journal=J%20Neurosci&volume=18&pages=7381-7393&publication_year=1998&author=Precht%2CJC&author=Emde%2CG&author=Wolfart%2CJ&author=Karamursel%2CS&author=Akoev%2CGN&author=Andrianov%2CYN&author=Bullock%2CTH"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR93">Quillet E, Krieg F, Dechamp N, Hervet C, Bérard A, Le Roy P, Guyomard R, Prunet P, Pottinger TG (2014) Quantitative trait loci for magnitude of the plasma cortisol response to confinement in rainbow trout. Genet, Anim. doi:<a href="https://doi.org/10.1111/age.12126" data-track="click_references" data-track-action="external reference" data-track-value="external reference" data-track-label="10.1111/age.12126">10.1111/age.12126</a> </p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 94" href="http://scholar.google.com/scholar_lookup?&title=Quantitative%20trait%20loci%20for%20magnitude%20of%20the%20plasma%20cortisol%20response%20to%20confinement%20in%20rainbow%20trout&publication_year=2014&author=Quillet%2CE&author=Krieg%2CF&author=Dechamp%2CN&author=Hervet%2CC&author=B%C3%A9rard%2CA&author=Roy%2CP&author=Guyomard%2CR&author=Prunet%2CP&author=Pottinger%2CTG"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR94">Railo H, Koivisto M (2012) Two means of suppressing visual awareness: a direct comparison of visual masking and transcranial magnetic stimulation. Cortex 48:333–343</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 95" href="http://scholar.google.com/scholar_lookup?&title=Two%20means%20of%20suppressing%20visual%20awareness%3A%20a%20direct%20comparison%20of%20visual%20masking%20and%20transcranial%20magnetic%20stimulation&journal=Cortex&volume=48&pages=333-343&publication_year=2012&author=Railo%2CH&author=Koivisto%2CM"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR95">Reilly SC, Quinn JP, Cossins AR, Sneddon LU (2008) Behavioural analysis of a nociceptive event in fish: comparisons between three species demonstrate specific responses. Appl Anim Behav Sci 114:248–259</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 96" href="http://scholar.google.com/scholar_lookup?&title=Behavioural%20analysis%20of%20a%20nociceptive%20event%20in%20fish%3A%20comparisons%20between%20three%20species%20demonstrate%20specific%20responses&journal=Appl%20Anim%20Behav%20Sci&volume=114&pages=248-259&publication_year=2008&author=Reilly%2CSC&author=Quinn%2CJP&author=Cossins%2CAR&author=Sneddon%2CLU"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR96">Roeser T, Naier H (2003) Visuomotor behaviors in larval zebrafish after GFP-guided laser ablation of the optic tectum. J Neurosci 23:3726–3734</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 97" href="http://scholar.google.com/scholar_lookup?&title=Visuomotor%20behaviors%20in%20larval%20zebrafish%20after%20GFP-guided%20laser%20ablation%20of%20the%20optic%20tectum&journal=J%20Neurosci&volume=23&pages=3726-3734&publication_year=2003&author=Roeser%2CT&author=Naier%2CH"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR97">Roques JAC, Abbink W, Geurds F, van de Vis H, Flik G (2010) Tailfin clipping, a painful procedure: studies on Nile tilapia and common carp. Physiol Behav 101:533–540</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 98" href="http://scholar.google.com/scholar_lookup?&title=Tailfin%20clipping%2C%20a%20painful%20procedure%3A%20studies%20on%20Nile%20tilapia%20and%20common%20carp&journal=Physiol%20Behav&volume=101&pages=533-540&publication_year=2010&author=Roques%2CJAC&author=Abbink%2CW&author=Geurds%2CF&author=Vis%2CH&author=Flik%2CG"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR98">Rose JD (2002) The neurobiological nature of fishes and the question of awareness and pain. Rev Fish Sci 10:1–38</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 99" href="http://scholar.google.com/scholar_lookup?&title=The%20neurobiological%20nature%20of%20fishes%20and%20the%20question%20of%20awareness%20and%20pain&journal=Rev%20Fish%20Sci&volume=10&pages=1-38&publication_year=2002&author=Rose%2CJD"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR99">Rose JD (2007) Anthropomorphism and ‘mental welfare’ of fishes. Dis Aquat Organ 75:139–154</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 100" href="http://scholar.google.com/scholar_lookup?&title=Anthropomorphism%20and%20%E2%80%98mental%20welfare%E2%80%99%20of%20fishes&journal=Dis%20Aquat%20Organ&volume=75&pages=139-154&publication_year=2007&author=Rose%2CJD"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR100">Rose JD, Arlinghaus R, Cooke SJ, Diggles BK, Sawynok W, Stevens ED, Wynne CDL (2014) Can fish really feel pain? Fish Fish 15:97–133</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 101" href="http://scholar.google.com/scholar_lookup?&title=Can%20fish%20really%20feel%20pain%3F&journal=Fish%20Fish&volume=15&pages=97-133&publication_year=2014&author=Rose%2CJD&author=Arlinghaus%2CR&author=Cooke%2CSJ&author=Diggles%2CBK&author=Sawynok%2CW&author=Stevens%2CED&author=Wynne%2CCDL"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR101">Saidel WM, Marquez-Houston K, Butler AB (2001) Identification of visual pallial telencephalon in the goldfish, <i>Carassius auratus</i>: a combined cytochrome oxidase and electrophysiological study. Brain Res 919:82–93</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 102" href="http://scholar.google.com/scholar_lookup?&title=Identification%20of%20visual%20pallial%20telencephalon%20in%20the%20goldfish%2C%20Carassius%20auratus%3A%20a%20combined%20cytochrome%20oxidase%20and%20electrophysiological%20study&journal=Brain%20Res&volume=919&pages=82-93&publication_year=2001&author=Saidel%2CWM&author=Marquez-Houston%2CK&author=Butler%2CAB"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR102">Saint-Amant L (2006) Development of motor networks in zebrafish embryos. Zebrafish 3:173–190</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 103" href="http://scholar.google.com/scholar_lookup?&title=Development%20of%20motor%20networks%20in%20zebrafish%20embryos&journal=Zebrafish&volume=3&pages=173-190&publication_year=2006&author=Saint-Amant%2CL"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR103">Savage GE (1968) Temporal factors in avoidance learning in normal and forebrainless goldfish (<i>Cartassius auratus</i>). Nature 218:1168–1169</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 104" href="http://scholar.google.com/scholar_lookup?&title=Temporal%20factors%20in%20avoidance%20learning%20in%20normal%20and%20forebrainless%20goldfish%20%28Cartassius%20auratus%29&journal=Nature&volume=218&pages=1168-1169&publication_year=1968&author=Savage%2CGE"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR104">Savage GE (1969) Some preliminary observations on the role of the telencephalon in food-reinforced behaviour in the goldfish, <i>Carassius auratus</i>. Anim Behav 17:760–772</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 105" href="http://scholar.google.com/scholar_lookup?&title=Some%20preliminary%20observations%20on%20the%20role%20of%20the%20telencephalon%20in%20food-reinforced%20behaviour%20in%20the%20goldfish%2C%20Carassius%20auratus&journal=Anim%20Behav&volume=17&pages=760-772&publication_year=1969&author=Savage%2CGE"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR105">Schubert D, Kotter R, Staiger JF (2007) Mapping functional connectivity in barrel-related columns reveals layer- and cell type-specific microcircuits. Brain Struct Funct 212:107–119</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 106" href="http://scholar.google.com/scholar_lookup?&title=Mapping%20functional%20connectivity%20in%20barrel-related%20columns%20reveals%20layer-%20and%20cell%20type-specific%20microcircuits&journal=Brain%20Struct%20Funct&volume=212&pages=107-119&publication_year=2007&author=Schubert%2CD&author=Kotter%2CR&author=Staiger%2CJF"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR106">Scott Weber E III (2011) Fish analgesia: pain, stress, fear aversion or nociception? Vet Clin Exot Anim 14:21–32</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 107" href="http://scholar.google.com/scholar_lookup?&title=Fish%20analgesia%3A%20pain%2C%20stress%2C%20fear%20aversion%20or%20nociception%3F&journal=Vet%20Clin%20Exot%20Anim&volume=14&pages=21-32&publication_year=2011&author=Scott%20Weber%2CE"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR107">Shimizu T (2009) Why can birds be so smart? Background, significance, and implications of the revised view of the avian brain. Comp Cogn Behav Rev 4:103–115</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 108" href="http://scholar.google.com/scholar_lookup?&title=Why%20can%20birds%20be%20so%20smart%3F%20Background%2C%20significance%2C%20and%20implications%20of%20the%20revised%20view%20of%20the%20avian%20brain&journal=Comp%20Cogn%20Behav%20Rev&volume=4&pages=103-115&publication_year=2009&author=Shimizu%2CT"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR108">Shimizu T, Bowers AN (1999) Visual circuits of the avian telencephalon. Behav Brain Res 98:183–191</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 109" href="http://scholar.google.com/scholar_lookup?&title=Visual%20circuits%20of%20the%20avian%20telencephalon&journal=Behav%20Brain%20Res&volume=98&pages=183-191&publication_year=1999&author=Shimizu%2CT&author=Bowers%2CAN"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR109">Shimizu T, Cox K, Karten HJ (1995) Intratelencephalic projections of the visual wulst in pigeons (<i>Columba livia</i>). J Comp Neurol 359:551–572</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 110" href="http://scholar.google.com/scholar_lookup?&title=Intratelencephalic%20projections%20of%20the%20visual%20wulst%20in%20pigeons%20%28Columba%20livia%29&journal=J%20Comp%20Neurol&volume=359&pages=551-572&publication_year=1995&author=Shimizu%2CT&author=Cox%2CK&author=Karten%2CHJ"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR110">Sillito AM, Cudeiro J, Jones HE (2006) Always returning: feedback and sensory processing in visual cortex and thalamus. TINS 29:307–316</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 111" href="http://scholar.google.com/scholar_lookup?&title=Always%20returning%3A%20feedback%20and%20sensory%20processing%20in%20visual%20cortex%20and%20thalamus&journal=TINS&volume=29&pages=307-316&publication_year=2006&author=Sillito%2CAM&author=Cudeiro%2CJ&author=Jones%2CHE"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR111">Sneddon LU (2003) The evidence for pain in fish: the use of morphine as an analgesic. Appl Anim Behave Sci 83:153–162</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 112" href="http://scholar.google.com/scholar_lookup?&title=The%20evidence%20for%20pain%20in%20fish%3A%20the%20use%20of%20morphine%20as%20an%20analgesic&journal=Appl%20Anim%20Behave%20Sci&volume=83&pages=153-162&publication_year=2003&author=Sneddon%2CLU"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR112">Sneddon LU (2004) Evolution of nociception in vertebrates: comparative analysis of lower vertebrates. Brain Res Rev 46:123–130</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 113" href="http://scholar.google.com/scholar_lookup?&title=Evolution%20of%20nociception%20in%20vertebrates%3A%20comparative%20analysis%20of%20lower%20vertebrates&journal=Brain%20Res%20Rev&volume=46&pages=123-130&publication_year=2004&author=Sneddon%2CLU"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR113">Sneddon LU (2011) Nociception or pain in fish. In: Farrell A (ed) Encyclopedia of fish physiology. Academic Press, London, pp 713–719. ISBN 978-0-12-374545-3</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 114" href="http://scholar.google.com/scholar_lookup?&title=Nociception%20or%20pain%20in%20fish&pages=713-719&publication_year=2011&author=Sneddon%2CLU"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR114">Sneddon LU, Braithwaite VA, Gentle MJ (2003) Do fishes have nociceptors? Evidence for the evolution of a vertebrate sensory system. Proc R Soc Lond B 270:1115–1121</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 115" href="http://scholar.google.com/scholar_lookup?&title=Do%20fishes%20have%20nociceptors%3F%20Evidence%20for%20the%20evolution%20of%20a%20vertebrate%20sensory%20system&journal=Proc%20R%20Soc%20Lond%20B&volume=270&pages=1115-1121&publication_year=2003&author=Sneddon%2CLU&author=Braithwaite%2CVA&author=Gentle%2CMJ"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR115">Soto-Moyano R, Galvez J, Vallejos C, Hernandez A (1988) Topical application of morphine to the rat somatosensory cortex produces analgesia to tonic pain. J Neurosci Res 19:511–514</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 116" href="http://scholar.google.com/scholar_lookup?&title=Topical%20application%20of%20morphine%20to%20the%20rat%20somatosensory%20cortex%20produces%20analgesia%20to%20tonic%20pain&journal=J%20Neurosci%20Res&volume=19&pages=511-514&publication_year=1988&author=Soto-Moyano%2CR&author=Galvez%2CJ&author=Vallejos%2CC&author=Hernandez%2CA"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR116">Takeuchi D, Hirabayashi T, Tamura K, Miyashita Y (2011) Reversal of interlaminar signal between sensory and memory processing in monkey temporal cortex. Science 331:1443–1447</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 117" href="http://scholar.google.com/scholar_lookup?&title=Reversal%20of%20interlaminar%20signal%20between%20sensory%20and%20memory%20processing%20in%20monkey%20temporal%20cortex&journal=Science&volume=331&pages=1443-1447&publication_year=2011&author=Takeuchi%2CD&author=Hirabayashi%2CT&author=Tamura%2CK&author=Miyashita%2CY"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR117">Taylor JJ, Borckardt JJ, Canterberyy M, Li X, Hanlon CA, Brown TR, George MS (2013) Naloxone-reversible modulation of pain circuitry by left prefrontal rTMS. Neuropsychopharmacology 38:1189–1197</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 118" href="http://scholar.google.com/scholar_lookup?&title=Naloxone-reversible%20modulation%20of%20pain%20circuitry%20by%20left%20prefrontal%20rTMS&journal=Neuropsychopharmacology&volume=38&pages=1189-1197&publication_year=2013&author=Taylor%2CJJ&author=Borckardt%2CJJ&author=Canterberyy%2CM&author=Li%2CX&author=Hanlon%2CCA&author=Brown%2CTR&author=George%2CMS"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR118">Thivierge J-P, Marcus GF (2007) The topographic brain: from neural connectivity to cognition. TINS 30:251–258</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 119" href="http://scholar.google.com/scholar_lookup?&title=The%20topographic%20brain%3A%20from%20neural%20connectivity%20to%20cognition&journal=TINS&volume=30&pages=251-258&publication_year=2007&author=Thivierge%2CJ-P&author=Marcus%2CGF"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR119">Tommerdahl M, Delemos KA, Vierck CJ, Favorov OV, Whitsel BL (1996) Anterior parietal cortical response to tactile and skin-heating stimuli applied to the same skin site. J Neurophysiol 75:2662–2670</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 120" href="http://scholar.google.com/scholar_lookup?&title=Anterior%20parietal%20cortical%20response%20to%20tactile%20and%20skin-heating%20stimuli%20applied%20to%20the%20same%20skin%20site&journal=J%20Neurophysiol&volume=75&pages=2662-2670&publication_year=1996&author=Tommerdahl%2CM&author=Delemos%2CKA&author=Vierck%2CCJ&author=Favorov%2COV&author=Whitsel%2CBL"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR120">Torquati K, Pizzella V, Babiloni C, del Gratta C, Della Penna S, Ferretti A, Franciotti R, Rossini PM, Romani GL (2005) Nociceptive and non-nociceptive sub-regions in the human somatosensory cortex: an MEG study using fMRI constraints. Neuroimage 26:48–56</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 121" href="http://scholar.google.com/scholar_lookup?&title=Nociceptive%20and%20non-nociceptive%20sub-regions%20in%20the%20human%20somatosensory%20cortex%3A%20an%20MEG%20study%20using%20fMRI%20constraints&journal=Neuroimage&volume=26&pages=48-56&publication_year=2005&author=Torquati%2CK&author=Pizzella%2CV&author=Babiloni%2CC&author=Gratta%2CC&author=Della%20Penna%2CS&author=Ferretti%2CA&author=Franciotti%2CR&author=Rossini%2CPM&author=Romani%2CGL"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR121">Tuoc TC, Radyushkin K, Tonchev AB, Piñon MC, Ashery-Padan R, Molnár Z, Davidoff MS, Stoykova A (2009) Selective cortical layering abnormalities and behavioral deficits in cortex-specific Pax6 knock-out mice. J Neurosci 29:8335–8349</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 122" href="http://scholar.google.com/scholar_lookup?&title=Selective%20cortical%20layering%20abnormalities%20and%20behavioral%20deficits%20in%20cortex-specific%20Pax6%20knock-out%20mice&journal=J%20Neurosci&volume=29&pages=8335-8349&publication_year=2009&author=Tuoc%2CTC&author=Radyushkin%2CK&author=Tonchev%2CAB&author=Pi%C3%B1on%2CMC&author=Ashery-Padan%2CR&author=Moln%C3%A1r%2CZ&author=Davidoff%2CMS&author=Stoykova%2CA"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR131">Vargas JP, Lopez JC, Portavella M (2009) What are the functions of fish brain pallium? Brain Res Bull 79:436–440</p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR122">Veit L, Nieder A (2013) Abstract rule neurons in the endbrain support intelligent behaviour in corvid songbirds. Nat Commun 4:2878. doi:<a href="https://doi.org/10.1038/ncomms3878" data-track="click_references" data-track-action="external reference" data-track-value="external reference" data-track-label="10.1038/ncomms3878">10.1038/ncomms3878</a> </p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 124" href="http://scholar.google.com/scholar_lookup?&title=Abstract%20rule%20neurons%20in%20the%20endbrain%20support%20intelligent%20behaviour%20in%20corvid%20songbirds&journal=Nat%20Commun&doi=10.1038%2Fncomms3878&volume=4&publication_year=2013&author=Veit%2CL&author=Nieder%2CA"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR123">Vierck CJ (2006) Animal models of pain. In: McMahon S, Koltzenburg M (eds) Wall and Melzack’s textbook of pain. Churchill, London</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 125" href="http://scholar.google.com/scholar_lookup?&title=Animal%20models%20of%20pain&publication_year=2006&author=Vierck%2CCJ"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR124">Watanabe S, Masuda S (2010) Integration of auditory and visual information in human face discrimination in pigeons. Behavioral and anatomical study. Behav Brain Res 207:61–69</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 126" href="http://scholar.google.com/scholar_lookup?&title=Integration%20of%20auditory%20and%20visual%20information%20in%20human%20face%20discrimination%20in%20pigeons.%20Behavioral%20and%20anatomical%20study&journal=Behav%20Brain%20Res&volume=207&pages=61-69&publication_year=2010&author=Watanabe%2CS&author=Masuda%2CS"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR125">Wolkers CPB, Junior BA, Menescal-de-Oliveira L, Hoffmann A (2013) Stress-induced antinociception in fish reversed by naloxone. PLoS ONE 8:e71175</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 127" href="http://scholar.google.com/scholar_lookup?&title=Stress-induced%20antinociception%20in%20fish%20reversed%20by%20naloxone&journal=PLoS%20ONE&volume=8&publication_year=2013&author=Wolkers%2CCPB&author=Junior%2CBA&author=Menescal-de-Oliveira%2CL&author=Hoffmann%2CA"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR126">Xie YF, Wang J, Huo FQ, Jia H, Tang JS (2004) μ but not δ and κ opioid receptor involvement in ventrolateral orbital cortex opioid-evoked antinociception in formalin test rats. Neuroscience 126:717–726</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 128" href="http://scholar.google.com/scholar_lookup?&title=%CE%BC%20but%20not%20%CE%B4%20and%20%CE%BA%20opioid%20receptor%20involvement%20in%20ventrolateral%20orbital%20cortex%20opioid-evoked%20antinociception%20in%20formalin%20test%20rats&journal=Neuroscience&volume=126&pages=717-726&publication_year=2004&author=Xie%2CYF&author=Wang%2CJ&author=Huo%2CFQ&author=Jia%2CH&author=Tang%2CJS"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR127">Yager D, Sharma SC, Grover BG (1977) Visual function in goldfish with unilateral and bilateral tectal ablation. Brain Res 137:267–275</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 129" href="http://scholar.google.com/scholar_lookup?&title=Visual%20function%20in%20goldfish%20with%20unilateral%20and%20bilateral%20tectal%20ablation&journal=Brain%20Res&volume=137&pages=267-275&publication_year=1977&author=Yager%2CD&author=Sharma%2CSC&author=Grover%2CBG"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR128">Yaksh TL, Rudy TA (1976) Analgesia mediated by a direct spinal action of narcotics. Science 192:1357–1358</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 130" href="http://scholar.google.com/scholar_lookup?&title=Analgesia%20mediated%20by%20a%20direct%20spinal%20action%20of%20narcotics&journal=Science&volume=192&pages=1357-1358&publication_year=1976&author=Yaksh%2CTL&author=Rudy%2CTA"> Google Scholar</a> </p></li><li class="c-article-references__item js-c-reading-companion-references-item"><p class="c-article-references__text" id="ref-CR129">Yamamoto N, Ito H (2008) Visual, lateral line, and auditory ascending pathways to the dorsal telencephalic area through the rostrolateral region of the lateral preglomerular nucleus in Cyprinids. J Comp Neurol 508:615–647</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 131" href="http://scholar.google.com/scholar_lookup?&title=Visual%2C%20lateral%20line%2C%20and%20auditory%20ascending%20pathways%20to%20the%20dorsal%20telencephalic%20area%20through%20the%20rostrolateral%20region%20of%20the%20lateral%20preglomerular%20nucleus%20in%20Cyprinids&journal=J%20Comp%20Neurol&volume=508&pages=615-647&publication_year=2008&author=Yamamoto%2CN&author=Ito%2CH"> Google Scholar</a> </p></li></ul><p class="c-article-references__download u-hide-print"><a data-track="click" data-track-action="download citation references" data-track-label="link" rel="nofollow" href="https://citation-needed.springer.com/v2/references/10.1007/s10539-014-9469-4?format=refman&flavour=references">Download references<svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-download-medium"></use></svg></a></p></div></div></div></section></div><section data-title="Conflict of interest"><div class="c-article-section" id="Ack1-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Ack1">Conflict of interest</h2><div class="c-article-section__content" id="Ack1-content"><p>The author states that he has not been paid for this work and has no conflict of interest.</p></div></div></section><section aria-labelledby="author-information" data-title="Author information"><div class="c-article-section" id="author-information-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="author-information">Author information</h2><div class="c-article-section__content" id="author-information-content"><h3 class="c-article__sub-heading" id="affiliations">Authors and Affiliations</h3><ol class="c-article-author-affiliation__list"><li id="Aff1"><p class="c-article-author-affiliation__address">School of Biomedical Sciences, University of Queensland, Brisbane, 4072, Australia</p><p class="c-article-author-affiliation__authors-list">Brian Key</p></li></ol><div class="u-js-hide u-hide-print" data-test="author-info"><span class="c-article__sub-heading">Authors</span><ol class="c-article-authors-search u-list-reset"><li id="auth-Brian-Key-Aff1"><span class="c-article-authors-search__title u-h3 js-search-name">Brian Key</span><div class="c-article-authors-search__list"><div class="c-article-authors-search__item c-article-authors-search__list-item--left"><a href="/search?dc.creator=Brian%20Key" class="c-article-button" data-track="click" data-track-action="author link - publication" data-track-label="link" rel="nofollow">View author publications</a></div><div class="c-article-authors-search__item c-article-authors-search__list-item--right"><p class="search-in-title-js c-article-authors-search__text">You can also search for this author in <span class="c-article-identifiers"><a class="c-article-identifiers__item" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=search&term=Brian%20Key" data-track="click" data-track-action="author link - pubmed" data-track-label="link" rel="nofollow">PubMed</a><span class="u-hide"> </span><a class="c-article-identifiers__item" href="http://scholar.google.co.uk/scholar?as_q=&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_sauthors=%22Brian%20Key%22&as_publication=&as_ylo=&as_yhi=&as_allsubj=all&hl=en" data-track="click" data-track-action="author link - scholar" data-track-label="link" rel="nofollow">Google Scholar</a></span></p></div></div></li></ol></div><h3 class="c-article__sub-heading" id="corresponding-author">Corresponding author</h3><p id="corresponding-author-list">Correspondence to <a id="corresp-c1" href="mailto:brian.key@uq.edu.au">Brian Key</a>.</p></div></div></section><section data-title="Rights and permissions"><div class="c-article-section" id="rightslink-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="rightslink">Rights and permissions</h2><div class="c-article-section__content" id="rightslink-content"> <p> <b>Open Access</b> This article is distributed under the terms of the Creative Commons Attribution License which permits any use, distribution, and reproduction in any medium, provided the original author(s) and the source are credited.</p> <p class="c-article-rights"><a data-track="click" data-track-action="view rights and permissions" data-track-label="link" href="https://s100.copyright.com/AppDispatchServlet?title=Fish%20do%20not%20feel%20pain%20and%20its%20implications%20for%20understanding%20phenomenal%20consciousness&author=Brian%20Key&contentID=10.1007%2Fs10539-014-9469-4&copyright=The%20Author%28s%29&publication=0169-3867&publicationDate=2014-12-16&publisherName=SpringerNature&orderBeanReset=true&oa=CC%20BY">Reprints and permissions</a></p></div></div></section><section aria-labelledby="article-info" data-title="About this article"><div class="c-article-section" id="article-info-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="article-info">About this article</h2><div class="c-article-section__content" id="article-info-content"><div class="c-bibliographic-information"><div class="u-hide-print c-bibliographic-information__column c-bibliographic-information__column--border"><a data-crossmark="10.1007/s10539-014-9469-4" target="_blank" rel="noopener" href="https://crossmark.crossref.org/dialog/?doi=10.1007/s10539-014-9469-4" data-track="click" data-track-action="Click Crossmark" data-track-label="link" data-test="crossmark"><img loading="lazy" width="57" height="81" alt="Check for updates. Verify currency and authenticity via CrossMark" src="data:image/svg+xml;base64,<svg height="81" width="57" xmlns="http://www.w3.org/2000/svg"><g fill="none" fill-rule="evenodd"><path d="m17.35 35.45 21.3-14.2v-17.03h-21.3" fill="#989898"/><path d="m38.65 35.45-21.3-14.2v-17.03h21.3" fill="#747474"/><path d="m28 .5c-12.98 0-23.5 10.52-23.5 23.5s10.52 23.5 23.5 23.5 23.5-10.52 23.5-23.5c0-6.23-2.48-12.21-6.88-16.62-4.41-4.4-10.39-6.88-16.62-6.88zm0 41.25c-9.8 0-17.75-7.95-17.75-17.75s7.95-17.75 17.75-17.75 17.75 7.95 17.75 17.75c0 4.71-1.87 9.22-5.2 12.55s-7.84 5.2-12.55 5.2z" fill="#535353"/><path d="m41 36c-5.81 6.23-15.23 7.45-22.43 2.9-7.21-4.55-10.16-13.57-7.03-21.5l-4.92-3.11c-4.95 10.7-1.19 23.42 8.78 29.71 9.97 6.3 23.07 4.22 30.6-4.86z" fill="#9c9c9c"/><path d="m.2 58.45c0-.75.11-1.42.33-2.01s.52-1.09.91-1.5c.38-.41.83-.73 1.34-.94.51-.22 1.06-.32 1.65-.32.56 0 1.06.11 1.51.35.44.23.81.5 1.1.81l-.91 1.01c-.24-.24-.49-.42-.75-.56-.27-.13-.58-.2-.93-.2-.39 0-.73.08-1.05.23-.31.16-.58.37-.81.66-.23.28-.41.63-.53 1.04-.13.41-.19.88-.19 1.39 0 1.04.23 1.86.68 2.46.45.59 1.06.88 1.84.88.41 0 .77-.07 1.07-.23s.59-.39.85-.68l.91 1c-.38.43-.8.76-1.28.99-.47.22-1 .34-1.58.34-.59 0-1.13-.1-1.64-.31-.5-.2-.94-.51-1.31-.91-.38-.4-.67-.9-.88-1.48-.22-.59-.33-1.26-.33-2.02zm8.4-5.33h1.61v2.54l-.05 1.33c.29-.27.61-.51.96-.72s.76-.31 1.24-.31c.73 0 1.27.23 1.61.71.33.47.5 1.14.5 2.02v4.31h-1.61v-4.1c0-.57-.08-.97-.25-1.21-.17-.23-.45-.35-.83-.35-.3 0-.56.08-.79.22-.23.15-.49.36-.78.64v4.8h-1.61zm7.37 6.45c0-.56.09-1.06.26-1.51.18-.45.42-.83.71-1.14.29-.3.63-.54 1.01-.71.39-.17.78-.25 1.18-.25.47 0 .88.08 1.23.24.36.16.65.38.89.67s.42.63.54 1.03c.12.41.18.84.18 1.32 0 .32-.02.57-.07.76h-4.36c.07.62.29 1.1.65 1.44.36.33.82.5 1.38.5.29 0 .57-.04.83-.13s.51-.21.76-.37l.55 1.01c-.33.21-.69.39-1.09.53-.41.14-.83.21-1.26.21-.48 0-.92-.08-1.34-.25-.41-.16-.76-.4-1.07-.7-.31-.31-.55-.69-.72-1.13-.18-.44-.26-.95-.26-1.52zm4.6-.62c0-.55-.11-.98-.34-1.28-.23-.31-.58-.47-1.06-.47-.41 0-.77.15-1.07.45-.31.29-.5.73-.58 1.3zm2.5.62c0-.57.09-1.08.28-1.53.18-.44.43-.82.75-1.13s.69-.54 1.1-.71c.42-.16.85-.24 1.31-.24.45 0 .84.08 1.17.23s.61.34.85.57l-.77 1.02c-.19-.16-.38-.28-.56-.37-.19-.09-.39-.14-.61-.14-.56 0-1.01.21-1.35.63-.35.41-.52.97-.52 1.67 0 .69.17 1.24.51 1.66.34.41.78.62 1.32.62.28 0 .54-.06.78-.17.24-.12.45-.26.64-.42l.67 1.03c-.33.29-.69.51-1.08.65-.39.15-.78.23-1.18.23-.46 0-.9-.08-1.31-.24-.4-.16-.75-.39-1.05-.7s-.53-.69-.7-1.13c-.17-.45-.25-.96-.25-1.53zm6.91-6.45h1.58v6.17h.05l2.54-3.16h1.77l-2.35 2.8 2.59 4.07h-1.75l-1.77-2.98-1.08 1.23v1.75h-1.58zm13.69 1.27c-.25-.11-.5-.17-.75-.17-.58 0-.87.39-.87 1.16v.75h1.34v1.27h-1.34v5.6h-1.61v-5.6h-.92v-1.2l.92-.07v-.72c0-.35.04-.68.13-.98.08-.31.21-.57.4-.79s.42-.39.71-.51c.28-.12.63-.18 1.04-.18.24 0 .48.02.69.07.22.05.41.1.57.17zm.48 5.18c0-.57.09-1.08.27-1.53.17-.44.41-.82.72-1.13.3-.31.65-.54 1.04-.71.39-.16.8-.24 1.23-.24s.84.08 1.24.24c.4.17.74.4 1.04.71s.54.69.72 1.13c.19.45.28.96.28 1.53s-.09 1.08-.28 1.53c-.18.44-.42.82-.72 1.13s-.64.54-1.04.7-.81.24-1.24.24-.84-.08-1.23-.24-.74-.39-1.04-.7c-.31-.31-.55-.69-.72-1.13-.18-.45-.27-.96-.27-1.53zm1.65 0c0 .69.14 1.24.43 1.66.28.41.68.62 1.18.62.51 0 .9-.21 1.19-.62.29-.42.44-.97.44-1.66 0-.7-.15-1.26-.44-1.67-.29-.42-.68-.63-1.19-.63-.5 0-.9.21-1.18.63-.29.41-.43.97-.43 1.67zm6.48-3.44h1.33l.12 1.21h.05c.24-.44.54-.79.88-1.02.35-.24.7-.36 1.07-.36.32 0 .59.05.78.14l-.28 1.4-.33-.09c-.11-.01-.23-.02-.38-.02-.27 0-.56.1-.86.31s-.55.58-.77 1.1v4.2h-1.61zm-47.87 15h1.61v4.1c0 .57.08.97.25 1.2.17.24.44.35.81.35.3 0 .57-.07.8-.22.22-.15.47-.39.73-.73v-4.7h1.61v6.87h-1.32l-.12-1.01h-.04c-.3.36-.63.64-.98.86-.35.21-.76.32-1.24.32-.73 0-1.27-.24-1.61-.71-.33-.47-.5-1.14-.5-2.02zm9.46 7.43v2.16h-1.61v-9.59h1.33l.12.72h.05c.29-.24.61-.45.97-.63.35-.17.72-.26 1.1-.26.43 0 .81.08 1.15.24.33.17.61.4.84.71.24.31.41.68.53 1.11.13.42.19.91.19 1.44 0 .59-.09 1.11-.25 1.57-.16.47-.38.85-.65 1.16-.27.32-.58.56-.94.73-.35.16-.72.25-1.1.25-.3 0-.6-.07-.9-.2s-.59-.31-.87-.56zm0-2.3c.26.22.5.37.73.45.24.09.46.13.66.13.46 0 .84-.2 1.15-.6.31-.39.46-.98.46-1.77 0-.69-.12-1.22-.35-1.61-.23-.38-.61-.57-1.13-.57-.49 0-.99.26-1.52.77zm5.87-1.69c0-.56.08-1.06.25-1.51.16-.45.37-.83.65-1.14.27-.3.58-.54.93-.71s.71-.25 1.08-.25c.39 0 .73.07 1 .2.27.14.54.32.81.55l-.06-1.1v-2.49h1.61v9.88h-1.33l-.11-.74h-.06c-.25.25-.54.46-.88.64-.33.18-.69.27-1.06.27-.87 0-1.56-.32-2.07-.95s-.76-1.51-.76-2.65zm1.67-.01c0 .74.13 1.31.4 1.7.26.38.65.58 1.15.58.51 0 .99-.26 1.44-.77v-3.21c-.24-.21-.48-.36-.7-.45-.23-.08-.46-.12-.7-.12-.45 0-.82.19-1.13.59-.31.39-.46.95-.46 1.68zm6.35 1.59c0-.73.32-1.3.97-1.71.64-.4 1.67-.68 3.08-.84 0-.17-.02-.34-.07-.51-.05-.16-.12-.3-.22-.43s-.22-.22-.38-.3c-.15-.06-.34-.1-.58-.1-.34 0-.68.07-1 .2s-.63.29-.93.47l-.59-1.08c.39-.24.81-.45 1.28-.63.47-.17.99-.26 1.54-.26.86 0 1.51.25 1.93.76s.63 1.25.63 2.21v4.07h-1.32l-.12-.76h-.05c-.3.27-.63.48-.98.66s-.73.27-1.14.27c-.61 0-1.1-.19-1.48-.56-.38-.36-.57-.85-.57-1.46zm1.57-.12c0 .3.09.53.27.67.19.14.42.21.71.21.28 0 .54-.07.77-.2s.48-.31.73-.56v-1.54c-.47.06-.86.13-1.18.23-.31.09-.57.19-.76.31s-.33.25-.41.4c-.09.15-.13.31-.13.48zm6.29-3.63h-.98v-1.2l1.06-.07.2-1.88h1.34v1.88h1.75v1.27h-1.75v3.28c0 .8.32 1.2.97 1.2.12 0 .24-.01.37-.04.12-.03.24-.07.34-.11l.28 1.19c-.19.06-.4.12-.64.17-.23.05-.49.08-.76.08-.4 0-.74-.06-1.02-.18-.27-.13-.49-.3-.67-.52-.17-.21-.3-.48-.37-.78-.08-.3-.12-.64-.12-1.01zm4.36 2.17c0-.56.09-1.06.27-1.51s.41-.83.71-1.14c.29-.3.63-.54 1.01-.71.39-.17.78-.25 1.18-.25.47 0 .88.08 1.23.24.36.16.65.38.89.67s.42.63.54 1.03c.12.41.18.84.18 1.32 0 .32-.02.57-.07.76h-4.37c.08.62.29 1.1.65 1.44.36.33.82.5 1.38.5.3 0 .58-.04.84-.13.25-.09.51-.21.76-.37l.54 1.01c-.32.21-.69.39-1.09.53s-.82.21-1.26.21c-.47 0-.92-.08-1.33-.25-.41-.16-.77-.4-1.08-.7-.3-.31-.54-.69-.72-1.13-.17-.44-.26-.95-.26-1.52zm4.61-.62c0-.55-.11-.98-.34-1.28-.23-.31-.58-.47-1.06-.47-.41 0-.77.15-1.08.45-.31.29-.5.73-.57 1.3zm3.01 2.23c.31.24.61.43.92.57.3.13.63.2.98.2.38 0 .65-.08.83-.23s.27-.35.27-.6c0-.14-.05-.26-.13-.37-.08-.1-.2-.2-.34-.28-.14-.09-.29-.16-.47-.23l-.53-.22c-.23-.09-.46-.18-.69-.3-.23-.11-.44-.24-.62-.4s-.33-.35-.45-.55c-.12-.21-.18-.46-.18-.75 0-.61.23-1.1.68-1.49.44-.38 1.06-.57 1.83-.57.48 0 .91.08 1.29.25s.71.36.99.57l-.74.98c-.24-.17-.49-.32-.73-.42-.25-.11-.51-.16-.78-.16-.35 0-.6.07-.76.21-.17.15-.25.33-.25.54 0 .14.04.26.12.36s.18.18.31.26c.14.07.29.14.46.21l.54.19c.23.09.47.18.7.29s.44.24.64.4c.19.16.34.35.46.58.11.23.17.5.17.82 0 .3-.06.58-.17.83-.12.26-.29.48-.51.68-.23.19-.51.34-.84.45-.34.11-.72.17-1.15.17-.48 0-.95-.09-1.41-.27-.46-.19-.86-.41-1.2-.68z" fill="#535353"/></g></svg>"></a></div><div class="c-bibliographic-information__column"><h3 class="c-article__sub-heading" id="citeas">Cite this article</h3><p class="c-bibliographic-information__citation">Key, B. Fish do not feel pain and its implications for understanding phenomenal consciousness. <i>Biol Philos</i> <b>30</b>, 149–165 (2015). https://doi.org/10.1007/s10539-014-9469-4</p><p class="c-bibliographic-information__download-citation u-hide-print"><a data-test="citation-link" data-track="click" data-track-action="download article citation" data-track-label="link" data-track-external="" rel="nofollow" href="https://citation-needed.springer.com/v2/references/10.1007/s10539-014-9469-4?format=refman&flavour=citation">Download citation<svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-download-medium"></use></svg></a></p><ul class="c-bibliographic-information__list" data-test="publication-history"><li class="c-bibliographic-information__list-item"><p>Received<span class="u-hide">: </span><span class="c-bibliographic-information__value"><time datetime="2014-04-14">14 April 2014</time></span></p></li><li class="c-bibliographic-information__list-item"><p>Accepted<span class="u-hide">: </span><span class="c-bibliographic-information__value"><time datetime="2014-12-06">06 December 2014</time></span></p></li><li class="c-bibliographic-information__list-item"><p>Published<span class="u-hide">: </span><span class="c-bibliographic-information__value"><time datetime="2014-12-16">16 December 2014</time></span></p></li><li class="c-bibliographic-information__list-item"><p>Issue Date<span class="u-hide">: </span><span class="c-bibliographic-information__value"><time datetime="2015-03">March 2015</time></span></p></li><li class="c-bibliographic-information__list-item c-bibliographic-information__list-item--full-width"><p><abbr title="Digital Object Identifier">DOI</abbr><span class="u-hide">: </span><span class="c-bibliographic-information__value">https://doi.org/10.1007/s10539-014-9469-4</span></p></li></ul><div data-component="share-box"><div class="c-article-share-box u-display-none" hidden=""><h3 class="c-article__sub-heading">Share this article</h3><p class="c-article-share-box__description">Anyone you share the following link with will be able to read this content:</p><button class="js-get-share-url c-article-share-box__button" type="button" id="get-share-url" data-track="click" data-track-label="button" data-track-external="" data-track-action="get shareable link">Get shareable link</button><div class="js-no-share-url-container u-display-none" hidden=""><p class="js-c-article-share-box__no-sharelink-info c-article-share-box__no-sharelink-info">Sorry, a shareable link is not currently available for this article.</p></div><div class="js-share-url-container u-display-none" hidden=""><p class="js-share-url c-article-share-box__only-read-input" id="share-url" data-track="click" data-track-label="button" data-track-action="select share url"></p><button class="js-copy-share-url c-article-share-box__button--link-like" type="button" id="copy-share-url" data-track="click" data-track-label="button" data-track-action="copy share url" data-track-external="">Copy to clipboard</button></div><p class="js-c-article-share-box__additional-info c-article-share-box__additional-info"> Provided by the Springer Nature SharedIt content-sharing initiative </p></div></div><h3 class="c-article__sub-heading">Keywords</h3><ul class="c-article-subject-list"><li class="c-article-subject-list__subject"><span><a href="/search?query=Fish&facet-discipline="Philosophy"" data-track="click" data-track-action="view keyword" data-track-label="link">Fish</a></span></li><li class="c-article-subject-list__subject"><span><a href="/search?query=Pain&facet-discipline="Philosophy"" data-track="click" data-track-action="view keyword" data-track-label="link">Pain</a></span></li><li class="c-article-subject-list__subject"><span><a href="/search?query=Phenomenal%20consciousness&facet-discipline="Philosophy"" data-track="click" data-track-action="view keyword" data-track-label="link">Phenomenal consciousness</a></span></li><li class="c-article-subject-list__subject"><span><a href="/search?query=Affective%20states&facet-discipline="Philosophy"" data-track="click" data-track-action="view keyword" data-track-label="link">Affective states</a></span></li><li class="c-article-subject-list__subject"><span><a href="/search?query=Avoidance%20learning&facet-discipline="Philosophy"" data-track="click" data-track-action="view keyword" data-track-label="link">Avoidance learning</a></span></li><li class="c-article-subject-list__subject"><span><a href="/search?query=Neocortex&facet-discipline="Philosophy"" data-track="click" data-track-action="view keyword" data-track-label="link">Neocortex</a></span></li><li class="c-article-subject-list__subject"><span><a href="/search?query=Pallium&facet-discipline="Philosophy"" data-track="click" data-track-action="view keyword" data-track-label="link">Pallium</a></span></li></ul><div data-component="article-info-list"></div></div></div></div></div></section> </div> </main> <div class="c-article-sidebar u-text-sm u-hide-print l-with-sidebar__sidebar" id="sidebar" data-container-type="reading-companion" data-track-component="reading companion"> <aside> <div class="app-card-service" data-test="article-checklist-banner"> <div> <a class="app-card-service__link" data-track="click_presubmission_checklist" data-track-context="article page top of reading companion" data-track-category="pre-submission-checklist" data-track-action="clicked article page checklist banner test 2 old version" data-track-label="link" href="https://beta.springernature.com/pre-submission?journalId=10539" data-test="article-checklist-banner-link"> <span class="app-card-service__link-text">Use our pre-submission checklist</span> <svg class="app-card-service__link-icon" aria-hidden="true" focusable="false"><use xlink:href="#icon-eds-i-arrow-right-small"></use></svg> </a> <p class="app-card-service__description">Avoid common mistakes on your manuscript.</p> </div> <div class="app-card-service__icon-container"> <svg class="app-card-service__icon" aria-hidden="true" focusable="false"> <use xlink:href="#icon-eds-i-clipboard-check-medium"></use> </svg> </div> </div> <div data-test="collections"> </div> <div data-test="editorial-summary"> </div> <div class="c-reading-companion"> <div class="c-reading-companion__sticky" data-component="reading-companion-sticky" data-test="reading-companion-sticky"> <div class="c-reading-companion__panel c-reading-companion__sections c-reading-companion__panel--active" id="tabpanel-sections"> <div class="u-lazy-ad-wrapper u-mt-16 u-hide" data-component-mpu><div class="c-ad c-ad--300x250"> <div class="c-ad__inner"> <p class="c-ad__label">Advertisement</p> <div id="div-gpt-ad-MPU1" class="div-gpt-ad grade-c-hide" data-pa11y-ignore data-gpt data-gpt-unitpath="/270604982/springerlink/10539/article" data-gpt-sizes="300x250" data-test="MPU1-ad" data-gpt-targeting="pos=MPU1;articleid=s10539-014-9469-4;"> </div> </div> </div> </div> </div> <div class="c-reading-companion__panel c-reading-companion__figures c-reading-companion__panel--full-width" id="tabpanel-figures"></div> <div class="c-reading-companion__panel c-reading-companion__references c-reading-companion__panel--full-width" id="tabpanel-references"></div> </div> </div> </aside> </div> </div> </article> <div class="app-elements"> <div class="eds-c-header__expander eds-c-header__expander--search" id="eds-c-header-popup-search"> <h2 class="eds-c-header__heading">Search</h2> <div class="u-container"> <search class="eds-c-header__search" role="search" aria-label="Search from the header"> <form method="GET" action="//link.springer.com/search" data-test="header-search" data-track="search" data-track-context="search from header" data-track-action="submit search form" data-track-category="unified header" data-track-label="form" > <label for="eds-c-header-search" class="eds-c-header__search-label">Search by keyword or author</label> <div class="eds-c-header__search-container"> <input id="eds-c-header-search" class="eds-c-header__search-input" autocomplete="off" name="query" type="search" value="" required> <button class="eds-c-header__search-button" type="submit"> <svg class="eds-c-header__icon" aria-hidden="true" focusable="false"> <use xlink:href="#icon-eds-i-search-medium"></use> </svg> <span class="u-visually-hidden">Search</span> </button> </div> </form> </search> </div> </div> <div class="eds-c-header__expander eds-c-header__expander--menu" id="eds-c-header-nav"> <h2 class="eds-c-header__heading">Navigation</h2> <ul class="eds-c-header__list"> <li class="eds-c-header__list-item"> <a class="eds-c-header__link" href="https://link.springer.com/journals/" data-track="nav_find_a_journal" data-track-context="unified header" data-track-action="click find a journal" data-track-category="unified header" data-track-label="link" > Find a journal </a> </li> <li class="eds-c-header__list-item"> <a class="eds-c-header__link" href="https://www.springernature.com/gp/authors" data-track="nav_how_to_publish" data-track-context="unified header" data-track-action="click publish with us link" data-track-category="unified header" data-track-label="link" > Publish with us </a> </li> <li class="eds-c-header__list-item"> <a class="eds-c-header__link" href="https://link.springernature.com/home/" data-track="nav_track_your_research" data-track-context="unified header" data-track-action="click track your research" data-track-category="unified header" data-track-label="link" > Track your research </a> </li> </ul> </div> <footer > <div class="eds-c-footer" > <div class="eds-c-footer__container"> <div class="eds-c-footer__grid eds-c-footer__group--separator"> <div class="eds-c-footer__group"> <h3 class="eds-c-footer__heading">Discover content</h3> <ul class="eds-c-footer__list"> <li class="eds-c-footer__item"><a class="eds-c-footer__link" href="https://link.springer.com/journals/a/1" data-track="nav_journals_a_z" data-track-action="journals a-z" data-track-context="unified footer" data-track-label="link">Journals A-Z</a></li> <li class="eds-c-footer__item"><a class="eds-c-footer__link" href="https://link.springer.com/books/a/1" data-track="nav_books_a_z" data-track-action="books a-z" data-track-context="unified footer" data-track-label="link">Books A-Z</a></li> </ul> </div> <div class="eds-c-footer__group"> <h3 class="eds-c-footer__heading">Publish with us</h3> <ul class="eds-c-footer__list"> <li class="eds-c-footer__item"><a class="eds-c-footer__link" href="https://link.springer.com/journals" data-track="nav_journal_finder" data-track-action="journal finder" data-track-context="unified footer" data-track-label="link">Journal finder</a></li> <li class="eds-c-footer__item"><a class="eds-c-footer__link" href="https://www.springernature.com/gp/authors" data-track="nav_publish_your_research" data-track-action="publish your research" data-track-context="unified footer" data-track-label="link">Publish your research</a></li> <li class="eds-c-footer__item"><a class="eds-c-footer__link" href="https://www.springernature.com/gp/open-research/about/the-fundamentals-of-open-access-and-open-research" data-track="nav_open_access_publishing" data-track-action="open access publishing" data-track-context="unified footer" data-track-label="link">Open access publishing</a></li> </ul> </div> <div class="eds-c-footer__group"> <h3 class="eds-c-footer__heading">Products and services</h3> <ul class="eds-c-footer__list"> <li class="eds-c-footer__item"><a class="eds-c-footer__link" href="https://www.springernature.com/gp/products" data-track="nav_our_products" data-track-action="our products" data-track-context="unified footer" data-track-label="link">Our products</a></li> <li class="eds-c-footer__item"><a class="eds-c-footer__link" href="https://www.springernature.com/gp/librarians" data-track="nav_librarians" data-track-action="librarians" data-track-context="unified footer" data-track-label="link">Librarians</a></li> <li class="eds-c-footer__item"><a class="eds-c-footer__link" href="https://www.springernature.com/gp/societies" data-track="nav_societies" data-track-action="societies" data-track-context="unified footer" data-track-label="link">Societies</a></li> <li class="eds-c-footer__item"><a class="eds-c-footer__link" href="https://www.springernature.com/gp/partners" data-track="nav_partners_and_advertisers" data-track-action="partners and advertisers" data-track-context="unified footer" data-track-label="link">Partners and advertisers</a></li> </ul> </div> <div class="eds-c-footer__group"> <h3 class="eds-c-footer__heading">Our imprints</h3> <ul class="eds-c-footer__list"> <li class="eds-c-footer__item"><a class="eds-c-footer__link" href="https://www.springer.com/" data-track="nav_imprint_Springer" data-track-action="Springer" data-track-context="unified footer" data-track-label="link">Springer</a></li> <li class="eds-c-footer__item"><a class="eds-c-footer__link" href="https://www.nature.com/" data-track="nav_imprint_Nature_Portfolio" data-track-action="Nature Portfolio" data-track-context="unified footer" data-track-label="link">Nature Portfolio</a></li> <li class="eds-c-footer__item"><a class="eds-c-footer__link" href="https://www.biomedcentral.com/" data-track="nav_imprint_BMC" data-track-action="BMC" data-track-context="unified footer" data-track-label="link">BMC</a></li> <li class="eds-c-footer__item"><a class="eds-c-footer__link" href="https://www.palgrave.com/" data-track="nav_imprint_Palgrave_Macmillan" data-track-action="Palgrave Macmillan" data-track-context="unified footer" data-track-label="link">Palgrave Macmillan</a></li> <li class="eds-c-footer__item"><a class="eds-c-footer__link" href="https://www.apress.com/" data-track="nav_imprint_Apress" data-track-action="Apress" data-track-context="unified footer" data-track-label="link">Apress</a></li> </ul> </div> </div> </div> <div class="eds-c-footer__container"> <nav aria-label="footer navigation"> <ul class="eds-c-footer__links"> <li class="eds-c-footer__item"> <button class="eds-c-footer__link" data-cc-action="preferences" data-track="dialog_manage_cookies" data-track-action="Manage cookies" data-track-context="unified footer" data-track-label="link"><span class="eds-c-footer__button-text">Your privacy choices/Manage cookies</span></button> </li> <li class="eds-c-footer__item"> <a class="eds-c-footer__link" href="https://www.springernature.com/gp/legal/ccpa" data-track="nav_california_privacy_statement" data-track-action="california privacy statement" data-track-context="unified footer" data-track-label="link">Your US state privacy rights</a> </li> <li class="eds-c-footer__item"> <a class="eds-c-footer__link" href="https://www.springernature.com/gp/info/accessibility" data-track="nav_accessibility_statement" data-track-action="accessibility statement" data-track-context="unified footer" data-track-label="link">Accessibility statement</a> </li> <li class="eds-c-footer__item"> <a class="eds-c-footer__link" href="https://link.springer.com/termsandconditions" data-track="nav_terms_and_conditions" data-track-action="terms and conditions" data-track-context="unified footer" data-track-label="link">Terms and conditions</a> </li> <li class="eds-c-footer__item"> <a class="eds-c-footer__link" href="https://link.springer.com/privacystatement" data-track="nav_privacy_policy" data-track-action="privacy policy" data-track-context="unified footer" data-track-label="link">Privacy policy</a> </li> <li class="eds-c-footer__item"> <a class="eds-c-footer__link" href="https://support.springernature.com/en/support/home" data-track="nav_help_and_support" data-track-action="help and support" data-track-context="unified footer" data-track-label="link">Help and support</a> </li> <li class="eds-c-footer__item"> <a class="eds-c-footer__link" href="https://support.springernature.com/en/support/solutions/articles/6000255911-subscription-cancellations" data-track-action="cancel contracts here">Cancel contracts here</a> </li> </ul> </nav> <div class="eds-c-footer__user"> <p class="eds-c-footer__user-info"> <span data-test="footer-user-ip">8.222.208.146</span> </p> <p class="eds-c-footer__user-info" data-test="footer-business-partners">Not affiliated</p> </div> <a href="https://www.springernature.com/" class="eds-c-footer__link"> <img src="/oscar-static/images/logo-springernature-white-19dd4ba190.svg" alt="Springer Nature" loading="lazy" width="200" height="20"/> </a> <p class="eds-c-footer__legal" data-test="copyright">© 2024 Springer Nature</p> </div> </div> </footer> </div> </body> </html>