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It is an old science, having its beginnings in prehistoric times.<ref>{{cite web |last=Rotimi |first=Booktionary |title=Anatomy |url=https://www.preps.ng/jamb-subject-combination-for-anatomy/ |url-status=live |archive-url=https://web.archive.org/web/20170801080440/https://www.booktionary.com.ng/index_files/Page1935.htm |archive-date=1 August 2017 |access-date=18 June 2017}}</ref> Anatomy is inherently tied to [[developmental biology]], [[embryology]], [[comparative anatomy]], [[evolutionary biology]], and [[phylogeny]],<ref name="intro HGray">{{cite web |last=Gray |first=Henry |year=1918 |title=Introduction |url=https://www.bartleby.com/107/1.html |archive-url=https://web.archive.org/web/20070316005206/https://www.bartleby.com/107/1.html |archive-date=16 March 2007 |access-date=19 March 2007 |website=Anatomy of the Human Body |via=[[Bartleby.com]] |edition=20th}}</ref> as these are the processes by which anatomy is generated, both over immediate and long-term timescales. Anatomy and [[physiology]], which study the structure and [[function (biology)|function]] of organisms and their parts respectively, make a natural pair of related disciplines, and are often studied together. Human anatomy is one of the essential [[basic sciences]] that are applied in medicine, and is often studied alongside [[physiology]].<ref>{{cite journal |author=Arráez-Aybar |display-authors=etal |year=2010 |title=Relevance of human anatomy in daily clinical practice |journal=[[Annals of Anatomy]] |volume=192 |issue=6|pages=341–48 |doi=10.1016/j.aanat.2010.05.002 |pmid=20591641 }}</ref> Anatomy is a complex and dynamic field that is constantly evolving as discoveries are made. In recent years, there has been a significant increase in the use of advanced imaging techniques, such as [[MRI]] and [[CT scan]]s, which allow for more detailed and accurate visualizations of the body's structures. The discipline of anatomy is divided into [[macroscopic]] and [[microscopic]] parts. [[Macroscopic anatomy]], or gross anatomy, is the examination of an animal's body parts using unaided eyesight. Gross anatomy also includes the branch of [[superficial anatomy]]. Microscopic anatomy involves the use of optical instruments in the study of the [[tissue (biology)|tissues]] of various structures, known as [[histology]], and also in the study of [[cell biology|cells]]. The [[history of anatomy]] is characterized by a progressive understanding of the functions of the organs and structures of the human body. Methods have also improved dramatically, advancing from the examination of animals by [[dissection]] of carcasses and [[cadaver]]s (corpses)<ref name=":0">{{Cite journal |last=Ghosh |first=Sanjib Kumar |date=2017-03-02 |title=Human cadaveric dissection: a historical account from ancient Greece to the modern era |journal=Anatomy & Cell Biology |volume=48 |issue=3 |pages=153–169 |doi=10.5115/acb.2015.48.3.153 |pmc=4582158 |pmid=26417475}}</ref> to 20th-century [[medical imaging]] techniques, including [[Radiography|X-ray]], [[Ultrasound imaging|ultrasound]], and [[magnetic resonance imaging]].<ref name=":1" /> == Etymology and definition == [[File:An écorché figure (life-size), lying prone on a table Wellcome L0020561.jpg|thumb|A dissected body, lying prone on a table, by [[Charles Landseer]]]] Derived from the Greek {{lang|grc|ἀνατομή}} ''anatomē'' "dissection" (from {{lang|grc|ἀνατέμνω}} ''anatémnō'' "I cut up, cut open" from ἀνά ''aná'' "up", and τέμνω ''témnō'' "I cut"),<ref>O.D.E. 2nd edition 2005</ref> anatomy is the scientific study of the structure of [[organism]]s including their systems, [[organs]] and [[tissue (biology)|tissues]]. It includes the appearance and position of the various parts, the materials from which they are composed, and their relationships with other parts. Anatomy is quite distinct from [[physiology]] and [[biochemistry]], which deal respectively with the functions of those parts and the chemical processes involved. For example, an anatomist is concerned with the shape, size, position, structure, blood supply and innervation of an organ such as the liver; while a physiologist is interested in the production of [[bile]], the role of the liver in nutrition and the regulation of bodily functions.<ref name=Everyman>{{cite book |title=Everyman's Encyclopedia: Anatomy |editor-last=Bozman |editor-first=E. F.|year=1967 |publisher=J. M. Dent & Sons |asin=B0066E44EC |page=272 }}</ref> The discipline of anatomy can be subdivided into a number of branches, including gross or [[macroscopic]] anatomy and [[microscopic]] anatomy.<ref>{{cite web |url=https://medical-dictionary.thefreedictionary.com/microscopic+anatomy |title=Anatomy |year=2007 |work=The Free Dictionary |publisher=Farlex |access-date=8 July 2013 |archive-date=15 November 2018 |archive-url=https://web.archive.org/web/20181115225224/https://medical-dictionary.thefreedictionary.com/microscopic+anatomy |url-status=live }}</ref> [[Gross anatomy]] is the study of structures large enough to be seen with the naked eye, and also includes [[superficial anatomy]] or surface anatomy, the study by sight of the external body features. [[Microscopic anatomy]] is the study of structures on a microscopic scale, along with [[histology]] (the study of tissues), and [[embryology]] (the study of an organism in its immature condition).<ref name="intro HGray" /> Regional anatomy is the study of the interrelationships of all of the structures in a specific body region, such as the abdomen. In contrast, systemic anatomy is the study of the structures that make up a discrete body system—that is, a group of structures that work together to perform a unique body function, such as the digestive system.<ref name="openstax">{{cite book |author=J. Gordon Betts |title=Anatomy & physiology |date=2013 |publisher=OpenStax |chapter=1.1 Overview of Anatomy and Physiology |location=Houston, Texas |isbn=978-1-947172-04-3 |url=https://openstax.org/books/anatomy-and-physiology/pages/1-1-overview-of-anatomy-and-physiology |access-date=14 May 2023 |archive-date=3 April 2023 |archive-url=https://web.archive.org/web/20230403194414/https://openstax.org/books/anatomy-and-physiology/pages/1-1-overview-of-anatomy-and-physiology |url-status=live }}</ref> Anatomy can be studied using both invasive and non-invasive methods with the goal of obtaining information about the structure and organization of organs and systems.<ref name="intro HGray" /> Methods used include [[dissection]], in which a body is opened and its organs studied, and [[endoscopy]], in which a [[video camera]]-equipped instrument is inserted through a small incision in the body wall and used to explore the internal organs and other structures. [[Angiography]] using [[X-ray]]s or [[magnetic resonance angiography]] are methods to visualize blood vessels.<ref>{{cite journal | title=Use of Angiography to Outline the Cardiovascular Anatomy of the Sand Crab Portunus pelagicus Linnaeus |vauthors=Gribble N, Reynolds K | journal=Journal of Crustacean Biology | year=1993 | volume=13 | issue=4 | pages=627–637 | doi=10.1163/193724093x00192 | jstor=1549093}}</ref><ref>{{cite journal | title=Characterization of the Renal Portal System of the Common Green Iguana (Iguana iguana) by Digital Subtraction Imaging |vauthors=Benson KG, Forrest L | journal=Journal of Zoo and Wildlife Medicine | year=1999 | volume=30 | issue=2 | pages=235–241|pmid=10484138 }}</ref><ref>{{cite web |url=https://www.hopkinsmedicine.org/healthlibrary/test_procedures/cardiovascular/magnetic_resonance_angiography_mra_135,14/ |title=Magnetic Resonance Angiography (MRA) |publisher=Johns Hopkins Medicine |access-date=29 April 2014 |archive-date=7 October 2017 |archive-url=https://web.archive.org/web/20171007124356/https://www.hopkinsmedicine.org/healthlibrary/test_procedures/cardiovascular/magnetic_resonance_angiography_mra_135,14 |url-status=live }}</ref><ref>{{cite web | url=https://www.nhs.uk/conditions/angiography/Pages/Introduction.aspx | title=Angiography | publisher=[[National Health Service]] | access-date=29 April 2014 | archive-date=7 September 2017 | archive-url=https://web.archive.org/web/20170907045854/https://www.nhs.uk/conditions/Angiography/Pages/Introduction.aspx | url-status=live }}</ref> The term "anatomy" is commonly taken to refer to [[human anatomy]]. However, substantially similar structures and tissues are found throughout the rest of the animal kingdom, and the term also includes the anatomy of other animals. The term ''zootomy'' is also sometimes used to specifically refer to non-human animals. The structure and tissues of plants are of a dissimilar nature and they are studied in [[plant anatomy]].<ref name=Everyman/> == Animal tissues == [[File:Anima cell notext.svg|right|thumb|Stylized cutaway diagram of an animal cell (with flagella)]] The [[Kingdom (biology)|kingdom]] [[Animalia]] contains [[multicellular organism]]s that are [[heterotroph]]ic and [[motile]] (although some have secondarily adopted a [[Sessility (zoology)|sessile]] lifestyle). Most animals have bodies differentiated into separate [[Tissue (biology)|tissues]] and these animals are also known as [[eumetazoa]]ns. They have an internal [[digestion|digestive]] chamber, with one or two openings; the [[gamete]]s are produced in multicellular sex organs, and the [[zygote]]s include a [[blastula]] stage in their [[Embryogenesis|embryonic development]]. Metazoans do not include the [[sponge]]s, which have undifferentiated cells.<ref name=Dorit549>{{cite book |title=Zoology |url=https://archive.org/details/zoology0000dori |url-access=registration |last1=Dorit |first1=R. L. |last2=Walker |first2=W. F. |last3=Barnes |first3=R. D. |year=1991 |publisher=Saunders College Publishing |isbn=978-0-03-030504-7 |pages=[https://archive.org/details/zoology0000dori/page/547 547–549] }}</ref> Unlike [[plant cell]]s, [[animal cells]] have neither a cell wall nor [[chloroplast]]s. Vacuoles, when present, are more in number and much smaller than those in the plant cell. The body tissues are composed of numerous types of cells, including those found in muscles, nerves and skin. Each typically has a cell membrane formed of [[phospholipid]]s, [[cytoplasm]] and a [[Cell nucleus|nucleus]]. All of the different cells of an animal are derived from the embryonic [[germ layer]]s. Those simpler invertebrates which are formed from two germ layers of ectoderm and endoderm are called [[diploblastic]] and the more developed animals whose structures and organs are formed from three germ layers are called [[triploblastic]].<ref name=Ruppert60>{{cite book |title=Invertebrate Zoology, 7th edition |last1=Ruppert |first1=Edward E. |last2=Fox |first2=Richard, S. |last3=Barnes |first3=Robert D. |year=2004 |publisher=[[Cengage Learning]] |isbn=978-81-315-0104-7 |pages=59–60 }}</ref> All of a triploblastic animal's tissues and organs are derived from the three germ layers of the embryo, the [[ectoderm]], [[mesoderm]] and [[endoderm]]. Animal tissues can be grouped into four basic types: [[connective tissue|connective]], [[epithelial]], [[muscle]] and [[nervous tissue]]. [[File:Cartilage polarised.jpg|thumb|Hyaline cartilage at high magnification ([[H&E stain]])]] ===Connective tissue=== [[Connective tissue]]s are fibrous and made up of cells scattered among inorganic material called the [[extracellular matrix]]. Often called [[fascia]] (from the Latin "fascia," meaning "band" or "bandage"), connective tissues give shape to organs and holds them in place. The main types are loose connective tissue, [[adipose tissue]], fibrous connective tissue, [[cartilage]] and bone. The extracellular matrix contains [[protein]]s, the chief and most abundant of which is [[collagen]]. Collagen plays a major part in organizing and maintaining tissues. The matrix can be modified to form a skeleton to support or protect the body. An [[exoskeleton]] is a thickened, rigid [[cuticle]] which is stiffened by [[mineralisation (biology)|mineralization]], as in [[crustacean]]s or by the cross-linking of its proteins as in [[insect]]s. An [[endoskeleton]] is internal and present in all developed animals, as well as in many of those less developed.<ref name="Ruppert60" /> ===Epithelium=== [[File:Normal gastric mucosa low mag.jpg|thumb|right|Gastric mucosa at low magnification ([[H&E stain]])]] [[Epithelial tissue]] is composed of closely packed cells, bound to each other by [[cell adhesion molecule]]s, with little intercellular space. Epithelial cells can be [[Squamous epithelial cell|squamous]] (flat), [[Simple cuboidal epithelium|cuboidal]] or [[Columnar epithelial cell|columnar]] and rest on a [[basal lamina]], the upper layer of the [[basement membrane]],<ref>{{cite book|last=Dorland's|title=Illustrated Medical Dictionary|date=2012|publisher=Elsevier Saunders|isbn=978-1-4160-6257-8|page=203}}</ref> the lower layer is the reticular lamina lying next to the connective tissue in the extracellular matrix secreted by the epithelial cells.<ref>{{cite book|last=Dorland's|title=Illustrated Medical Dictionary|date=2012|publisher=Elsevier Saunders|isbn=978-1-4160-6257-8|page=1002}}</ref> There are many different types of epithelium, modified to suit a particular function. In the [[respiratory tract]] there is a type of [[pseudostratified ciliated columnar epithelium|ciliated]] epithelial lining; in the small intestine there are [[microvilli]] on the epithelial lining and in the large intestine there are [[intestinal villi]]. Skin consists of an outer layer of [[keratin]]ized stratified squamous epithelium that covers the exterior of the vertebrate body. [[Keratinocyte]]s make up to 95% of the cells in the [[epidermis (skin)|skin]].<ref>McGrath, J.A.; Eady, R.A.; Pope, F.M. (2004). Rook's Textbook of Dermatology (7th ed.). Blackwell Publishing. pp. 3.1–3.6. {{ISBN|978-0-632-06429-8}}.</ref> The epithelial cells on the external surface of the body typically secrete an extracellular matrix in the form of a [[cuticle]]. In simple animals this may just be a coat of [[glycoproteins]].<ref name="Ruppert60" /> In more advanced animals, many [[gland]]s are formed of epithelial cells.<ref>{{cite web |url=https://www.bio.davidson.edu/people/kabernd/BerndCV/Lab/EpithelialInfoWeb/Glandular%20Epithelium.html |title=Glandular epithelium |last=Bernd |first=Karen |year=2010 |work=Epithelial Cells |publisher=Davidson College |access-date=25 June 2013 |archive-url=https://web.archive.org/web/20200128053412/https://www.bio.davidson.edu/people/kabernd/BerndCV/Lab/EpithelialInfoWeb/Glandular%20Epithelium.html |archive-date=28 January 2020 |url-status=live }}</ref> ===Muscle tissue=== [[File:Skeletal muscle - cross section, nerve bundle.jpg|right|thumb|Cross section through [[skeletal muscle]] and a small [[nerve]] at high magnification ([[H&E stain]])]] [[Muscle cells]] (myocytes) form the active contractile tissue of the body. [[Muscle tissue]] functions to produce force and cause motion, either locomotion or movement within internal organs. Muscle is formed of contractile [[Myofibril|filaments]] and is separated into three main types; [[smooth muscle]], [[skeletal muscle]] and [[cardiac muscle]]. Smooth muscle has no [[Striated muscle tissue|striations]] when examined microscopically. It contracts slowly but maintains contractibility over a wide range of stretch lengths. It is found in such organs as [[sea anemone]] tentacles and the body wall of [[sea cucumber]]s. Skeletal muscle contracts rapidly but has a limited range of extension. It is found in the movement of appendages and jaws. Obliquely striated muscle is intermediate between the other two. The filaments are staggered and this is the type of muscle found in [[earthworm]]s that can extend slowly or make rapid contractions.<ref name=Ruppert103>{{cite book |title=Invertebrate Zoology, 7th edition |last1=Ruppert |first1=Edward E. |last2=Fox |first2=Richard, S. |last3=Barnes |first3=Robert D. |year=2004 |publisher=Cengage Learning |isbn=978-81-315-0104-7 |page=103 }}</ref> In higher animals striated muscles occur in bundles attached to bone to provide movement and are often arranged in antagonistic sets. Smooth muscle is found in the walls of the [[uterus]], [[bladder]], [[intestines]], [[stomach]], [[oesophagus]], [[respiratory airways]], and [[blood vessel]]s. [[Cardiac muscle]] is found only in the [[heart]], allowing it to contract and pump blood round the body. ===Nervous tissue=== {{See also|Neuroanatomy}} [[Nervous tissue]] is composed of many nerve cells known as [[neuron]]s which transmit information. In some slow-moving [[radially symmetrical]] marine animals such as [[ctenophore]]s and [[cnidarian]]s (including [[sea anemone]]s and [[jellyfish]]), the nerves form a [[nerve net]], but in most animals they are organized longitudinally into bundles. In simple animals, receptor neurons in the body wall cause a local reaction to a stimulus. In more complex animals, specialized receptor cells such as [[chemoreceptor]]s and [[photoreceptor cell|photoreceptors]] are found in groups and send messages along [[biological neural network|neural networks]] to other parts of the organism. Neurons can be connected together in [[ganglia]].<ref name=Ruppert104>{{cite book |title=Invertebrate Zoology, 7th edition |last1=Ruppert |first1=Edward E. |last2=Fox |first2=Richard, S. |last3=Barnes |first3=Robert D. |year=2004 |publisher=Cengage Learning |isbn=978-81-315-0104-7 |page=104 }}</ref> In higher animals, specialized receptors are the basis of sense organs and there is a [[central nervous system]] (brain and spinal cord) and a [[peripheral nervous system]]. The latter consists of [[Sensory neuron|sensory nerves]] that transmit information from sense organs and [[motor nerves]] that influence target organs.<ref>{{cite book|title=Grey's Anatomy: Descriptive and Applied |year=1944 |edition=28 |page=1038 |publisher=Langmans |editor1-last=Johnston | editor1-first= T.B |editor2-last=Whillis | editor2-first=J }}</ref><ref name=Ruppert107>{{cite book |title=Invertebrate Zoology, 7th edition |last1=Ruppert |first1=Edward E. |last2=Fox |first2=Richard, S. |last3=Barnes |first3=Robert D. |year=2004 |publisher=Cengage Learning |isbn=978-81-315-0104-7 |pages=105–107 }}</ref> The peripheral nervous system is divided into the [[somatic nervous system]] which conveys sensation and controls [[voluntary muscle]], and the [[autonomic nervous system]] which involuntarily controls [[smooth muscle]], certain glands and internal organs, including the [[stomach]].<ref>{{cite web | url=https://www.inkling.com/read/essential-clinical-anatomy-keith-moore-4th/introduction-to-clinical-anatomy/nervous-system | title=Essesntial Clinical Anatomy | publisher=Inkling | work=Nervous System | date=2010 | edition=4th | access-date=30 April 2014 | author1=Moore, K. | author2=Agur, A. | author3=Dalley, A. F. | archive-date=8 March 2021 | archive-url=https://web.archive.org/web/20210308131326/https://www.inkling.com/read/essential-clinical-anatomy-keith-moore-4th/introduction-to-clinical-anatomy/nervous-system | url-status=live }}</ref> == Vertebrate anatomy == {{see also|Vertebrate#Anatomy and morphology|Comparative anatomy}} [[File:VolRenderShearWarp.gif|thumb|upright|[[Mouse]] skull. The neck and most of the forelimbs are also seen.]] All [[vertebrate]]s have a similar basic [[body plan]] and at some point in their lives, mostly in the [[embryogenesis|embryonic]] stage, share the major [[chordate]] characteristics: a stiffening rod, the [[notochord]]; a dorsal hollow tube of nervous material, the [[neural tube]]; [[pharyngeal arch]]es; and a tail posterior to the anus. The [[spinal cord]] is protected by the [[vertebral column]] and is above the notochord, and the [[gastrointestinal tract]] is below it.<ref>{{cite web |last=Waggoner |first=Ben |title=Vertebrates: More on Morphology |url=https://www.ucmp.berkeley.edu/vertebrates/vertmm.html |publisher=UCMP |access-date=13 July 2011 |archive-date=10 October 2018 |archive-url=https://web.archive.org/web/20181010104933/https://www.ucmp.berkeley.edu/vertebrates/vertmm.html |url-status=dead }}</ref> Nervous tissue is derived from the [[ectoderm]], connective tissues are derived from [[mesoderm]], and gut is derived from the [[endoderm]]. At the posterior end is a tail which continues the spinal cord and vertebrae but not the gut. The mouth is found at the anterior end of the animal, and the [[anus]] at the base of the tail.<ref>{{cite book |title=The Vertebrate Body |last=Romer |first=Alfred Sherwood |year=1985 |publisher=Holt Rinehart & Winston |isbn=978-0-03-058446-6 }}</ref> The defining characteristic of a vertebrate is the [[vertebral column]], formed in the development of the segmented series of [[vertebra]]e. In most vertebrates the notochord becomes the [[nucleus pulposus]] of the [[intervertebral disc]]s. However, a few vertebrates, such as the [[sturgeon]] and the [[coelacanth]], retain the notochord into adulthood.<ref>{{cite book|title=Functional anatomy of the vertebrates: an evolutionary perspective|year=2001|publisher=Harcourt College Publishers|isbn=978-0-03-022369-3|author=Liem, Karel F.|author2=Warren Franklin Walker|page=277}}</ref> [[Jawed vertebrates]] are typified by paired appendages, fins or legs, which may be secondarily lost. The limbs of vertebrates are considered to be [[Homology (biology)|homologous]] because the same underlying skeletal structure was inherited from their last common ancestor. This is one of the arguments put forward by [[Charles Darwin]] to support his theory of [[evolution]].<ref>{{cite web |url=https://ncse.com/evolution/science/what-is-homology |title=What is Homology? |date=17 October 2008 |publisher=National Center for Science Education |access-date=28 June 2013 |archive-date=31 March 2019 |archive-url=https://web.archive.org/web/20190331065618/https://ncse.com/evolution/science/what-is-homology |url-status=live }}</ref> === Fish anatomy === {{main|Fish anatomy}} [[File:Internal anatomy of a fish (cutaway diagram).jpg|thumb|left|Cutaway diagram showing various organs of a fish]] The body of a [[fish]] is divided into a head, trunk and tail, although the divisions between the three are not always externally visible. The skeleton, which forms the support structure inside the fish, is either made of cartilage, in [[cartilaginous fish]], or bone in [[bony fish]]. The main skeletal element is the vertebral column, composed of articulating [[vertebra]]e which are lightweight yet strong. The ribs attach to the spine and there are no [[Limb (anatomy)|limbs]] or limb girdles. The main external features of the fish, the [[fish fin|fins]], are composed of either bony or soft spines called rays, which with the exception of the [[caudal fin]]s, have no direct connection with the spine. They are supported by the muscles which compose the main part of the trunk.<ref name=Dorit818/> The heart has two chambers and pumps the blood through the respiratory surfaces of the [[gill]]s and on round the body in a single circulatory loop.<ref>{{cite web|url=https://library.thinkquest.org/C003758/Development/fish.htm |title=The fish heart |work=ThinkQuest |publisher=Oracle |access-date=27 June 2013 |url-status=dead |archive-url=https://web.archive.org/web/20120428001536/https://library.thinkquest.org/C003758/Development/fish.htm |archive-date=28 April 2012 }}</ref> The eyes are adapted for seeing underwater and have only local vision. There is an inner ear but no external or [[middle ear]]. Low frequency vibrations are detected by the [[lateral line]] system of sense organs that run along the length of the sides of fish, and these respond to nearby movements and to changes in water pressure.<ref name=Dorit818>{{cite book |title=Zoology |url=https://archive.org/details/zoology0000dori |url-access=registration |last1=Dorit |first1=R. L. |last2=Walker |first2=W. F. |last3=Barnes |first3=R. D. |year=1991 |publisher=Saunders College Publishing |isbn=978-0-03-030504-7 |pages=[https://archive.org/details/zoology0000dori/page/816 816–818] }}</ref> Sharks and rays are [[Basal (phylogenetics)|basal]] fish with numerous [[Primitive (phylogenetics)|primitive]] anatomical features similar to those of ancient fish, including skeletons composed of cartilage. Their bodies tend to be dorso-ventrally flattened, they usually have five pairs of gill slits and a large mouth set on the underside of the head. The dermis is covered with separate dermal [[placoid scales]]. They have a [[cloaca]] into which the urinary and genital passages open, but not a [[swim bladder]]. Cartilaginous fish produce a small number of large, [[Egg yolk|yolky]] eggs. Some species are [[ovoviviparous]] and the young develop internally but others are [[oviparous]] and the larvae develop externally in egg cases.<ref name=Kotpal>{{cite book |title=Modern Text Book of Zoology: Vertebrates |last=Kotpal |first=R. L. |year=2010 |publisher=Rastogi Publications |isbn=978-81-7133-891-7 |page=193 |url=https://books.google.com/books?id=U7N1j-8LMsEC&pg=PA199 }}</ref> The bony fish lineage shows more [[Derived trait|derived]] anatomical traits, often with major evolutionary changes from the features of ancient fish. They have a bony skeleton, are generally laterally flattened, have five pairs of gills protected by an [[operculum (fish)|operculum]], and a mouth at or near the tip of the snout. The dermis is covered with overlapping [[Fish scale|scales]]. Bony fish have a swim bladder which helps them maintain a constant depth in the water column, but not a cloaca. They mostly [[Spawn (biology)|spawn]] a large number of small eggs with little yolk which they broadcast into the water column.<ref name=Kotpal/> === Amphibian anatomy === {{main|Amphibian anatomy}} [[File:Ceratophrys cornuta skeleton front.jpg|thumb|left|alt=Frog skeleton|Skeleton of [[Surinam horned frog]] (''Ceratophrys cornuta'')]] [[File:Frog anatomy tags.PNG|thumb|upright|Plastic model of a frog]] [[Amphibian]]s are a [[Class (biology)|class]] of animals comprising [[frog]]s, [[salamander]]s and [[caecilian]]s. They are [[tetrapod]]s, but the caecilians and a few species of salamander have either no limbs or their limbs are much reduced in size. Their main bones are hollow and lightweight and are fully ossified and the vertebrae interlock with each other and have [[articular processes]]. Their ribs are usually short and may be fused to the vertebrae. Their skulls are mostly broad and short, and are often incompletely ossified. Their skin contains little [[keratin]] and lacks scales, but contains many [[mucous gland]]s and in some species, poison glands. The hearts of amphibians have three chambers, two [[atrium (heart)|atria]] and one [[ventricle (heart)|ventricle]]. They have a [[urinary bladder]] and [[metabolic waste#nitrogen wastes|nitrogenous waste products]] are excreted primarily as [[urea]]. Amphibians breathe by means of [[buccal pumping]], a pump action in which air is first drawn into the [[Buccopharyngeal membrane|buccopharyngeal]] region through the nostrils. These are then closed and the air is forced into the lungs by contraction of the throat.<ref>{{cite book |title=A Natural History of Amphibians |last1=Stebbins |first1=Robert C. |author-link1=Robert C. Stebbins|last2=Cohen |first2=Nathan W. |year=1995 |publisher=Princeton University Press |isbn=978-0-691-03281-8 |pages=24–25 }}</ref> They supplement this with [[gas exchange]] through the skin which needs to be kept moist.<ref name=Dorit859>{{cite book |title=Zoology |url=https://archive.org/details/zoology0000dori/page/843 |url-access=registration |last1=Dorit |first1=R. L. |last2=Walker |first2=W. F. |last3=Barnes |first3=R. D. |year=1991 |publisher=Saunders College Publishing |isbn=978-0-03-030504-7 |pages=[https://archive.org/details/zoology0000dori/page/843 843–859] }}</ref> In frogs the pelvic girdle is robust and the hind legs are much longer and stronger than the forelimbs. The feet have four or five digits and the toes are often webbed for swimming or have suction pads for climbing. Frogs have large eyes and no tail. Salamanders resemble lizards in appearance; their short legs project sideways, the belly is close to or in contact with the ground and they have a long tail. Caecilians superficially resemble [[earthworm]]s and are limbless. They burrow by means of zones of muscle contractions which move along the body and they swim by undulating their body from side to side.<ref>{{cite book |title=A Natural History of Amphibians |last1=Stebbins |first1=Robert C. |author-link1=Robert C. Stebbins|last2=Cohen |first2=Nathan W. |year=1995 |publisher=Princeton University Press |isbn=978-0-691-03281-8 |pages=26–35 }}</ref> === Reptile anatomy === {{main|Reptile anatomy}} [[File:Crotalus atrox -Museum of Osteology, Oklahoma City, Oklahoma, USA-29Aug2012.jpg|thumb|left|Skeleton of a [[western diamondback rattlesnake]]]] [[Reptile]]s are a class of [[animal]]s comprising [[turtle]]s, [[tuatara]]s, [[lizard]]s, [[snake]]s and [[crocodile]]s. They are [[tetrapod]]s, but the snakes and a few species of lizard either have no limbs or their limbs are much reduced in size. Their bones are better ossified and their skeletons stronger than those of amphibians. The teeth are conical and mostly uniform in size. The surface cells of the epidermis are modified into horny scales which create a waterproof layer. Reptiles are unable to use their skin for respiration as do amphibians and have a more efficient respiratory system drawing air into their [[lung]]s by expanding their chest walls. The heart resembles that of the amphibian but there is a septum which more completely separates the oxygenated and deoxygenated bloodstreams. The reproductive system has evolved for internal fertilization, with a [[copulatory organ]] present in most species. The eggs are surrounded by [[Amniote|amniotic membranes]] which prevents them from drying out and are laid on land, or [[Ovoviviparity|develop internally]] in some species. The bladder is small as nitrogenous waste is excreted as [[uric acid]].<ref name=Dorit865>{{cite book |title=Zoology |url=https://archive.org/details/zoology0000dori/page/861 |url-access=registration |last1=Dorit |first1=R. L. |last2=Walker |first2=W. F. |last3=Barnes |first3=R. D. |year=1991 |publisher=Saunders College Publishing |isbn=978-0-03-030504-7 |pages=[https://archive.org/details/zoology0000dori/page/861 861–865] }}</ref> Turtles are notable for their protective shells. They have an inflexible trunk encased in a horny [[carapace]] above and a [[plastron]] below. These are formed from bony plates embedded in the dermis which are overlain by horny ones and are partially fused with the ribs and spine. The neck is long and flexible and the head and the legs can be drawn back inside the shell. Turtles are vegetarians and the typical reptile teeth have been replaced by sharp, horny plates. In aquatic species, the front legs are modified into flippers.<ref name=Dorit868>{{cite book |title=Zoology |url=https://archive.org/details/zoology0000dori/page/865 |url-access=registration |last1=Dorit |first1=R. L. |last2=Walker |first2=W. F. |last3=Barnes |first3=R. D. |year=1991 |publisher=Saunders College Publishing |isbn=978-0-03-030504-7 |pages=[https://archive.org/details/zoology0000dori/page/865 865–868] }}</ref> '''Tuataras''' superficially resemble lizards but the lineages diverged in the [[Triassic]] period. There is one living species, ''[[Sphenodon punctatus]]''. The skull has two openings (fenestrae) on either side and the jaw is rigidly attached to the skull. There is one row of teeth in the lower jaw and this fits between the two rows in the upper jaw when the animal chews. The teeth are merely projections of bony material from the jaw and eventually wear down. The brain and heart are more primitive than those of other reptiles, and the lungs have a single chamber and lack [[bronchi]]. The tuatara has a well-developed [[parietal eye]] on its forehead.<ref name=Dorit868/> Lizards have skulls with only one [[Nasal fenestra|fenestra]] on each side, the lower bar of bone below the second fenestra having been lost. This results in the jaws being less rigidly attached which allows the mouth to open wider. Lizards are mostly quadrupeds, with the trunk held off the ground by short, sideways-facing legs, but a few species have no limbs and resemble snakes. Lizards have moveable eyelids, eardrums are present and some species have a central parietal eye.<ref name=Dorit868/> Snakes are closely related to lizards, having branched off from a common ancestral lineage during the [[Cretaceous]] period, and they share many of the same features. The skeleton consists of a skull, a hyoid bone, spine and ribs though a few species retain a vestige of the pelvis and rear limbs in the form of [[pelvic spur]]s. The bar under the second fenestra has also been lost and the jaws have extreme flexibility allowing the snake to swallow its prey whole. Snakes lack moveable eyelids, the eyes being covered by transparent "spectacle" scales. They do not have eardrums but can detect ground vibrations through the bones of their skull. Their forked tongues are used as organs of taste and smell and some species have sensory pits on their heads enabling them to locate warm-blooded prey.<ref name=Dorit870>{{cite book |title=Zoology |url=https://archive.org/details/zoology0000dori/page/870 |url-access=registration |last1=Dorit |first1=R. L. |last2=Walker |first2=W. F. |last3=Barnes |first3=R. D. |year=1991 |publisher=Saunders College Publishing |isbn=978-0-03-030504-7 |page=[https://archive.org/details/zoology0000dori/page/870 870] }}</ref> [[Crocodilians]] are large, low-slung aquatic reptiles with long snouts and large numbers of teeth. The head and trunk are dorso-ventrally flattened and the tail is laterally compressed. It undulates from side to side to force the animal through the water when swimming. The tough keratinized scales provide body armour and some are fused to the skull. The nostrils, eyes and ears are elevated above the top of the flat head enabling them to remain above the surface of the water when the animal is floating. Valves seal the nostrils and ears when it is submerged. Unlike other reptiles, crocodilians have hearts with four chambers allowing complete separation of oxygenated and deoxygenated blood.<ref name=Dorit874>{{cite book |title=Zoology |url=https://archive.org/details/zoology0000dori/page/874 |url-access=registration |last1=Dorit |first1=R. L. |last2=Walker |first2=W. F. |last3=Barnes |first3=R. D. |year=1991 |publisher=Saunders College Publishing |isbn=978-0-03-030504-7 |page=[https://archive.org/details/zoology0000dori/page/874 874] }}</ref> === Bird anatomy === {{main|Bird anatomy}} [[File:Duerer wing of a blue roller.jpg|thumb|Part of a wing. [[Albrecht Dürer]], {{circa|1500–1512}}]] [[Bird]]s are [[tetrapod]]s but though their hind limbs are used for walking or hopping, their front limbs are [[wing]]s covered with [[feather]]s and adapted for flight. Birds are [[endotherm]]ic, have a high [[metabolic rate]], a light [[skeletal system]] and powerful [[muscle]]s. The long bones are thin, hollow and very light. Air sac extensions from the lungs occupy the centre of some bones. The sternum is wide and usually has a keel and the caudal vertebrae are fused. There are no teeth and the narrow jaws are adapted into a horn-covered beak. The eyes are relatively large, particularly in nocturnal species such as owls. They face forwards in predators and sideways in ducks.<ref name=Dorit895>{{cite book |title=Zoology |url=https://archive.org/details/zoology0000dori/page/881 |url-access=registration |last1=Dorit |first1=R. L. |last2=Walker |first2=W. F. |last3=Barnes |first3=R. D. |year=1991 |publisher=Saunders College Publishing |isbn=978-0-03-030504-7 |pages=[https://archive.org/details/zoology0000dori/page/881 881–895] }}</ref> The feathers are outgrowths of the [[epidermis (zoology)|epidermis]] and are found in localized bands from where they fan out over the skin. Large flight feathers are found on the wings and tail, contour feathers cover the bird's surface and fine down occurs on young birds and under the contour feathers of water birds. The only cutaneous gland is the single [[uropygial gland]] near the base of the tail. This produces an oily secretion that waterproofs the feathers when the bird [[personal grooming|preens]]. There are scales on the legs, feet and claws on the tips of the toes.<ref name=Dorit895/> === Mammal anatomy === {{Main|Mammal anatomy}} [[Mammal]]s are a diverse class of animals, mostly terrestrial but some are aquatic and others have evolved flapping or gliding flight. They mostly have four limbs, but some aquatic mammals have no limbs or limbs modified into fins, and the forelimbs of bats are modified into wings. The legs of most mammals are situated below the trunk, which is held well clear of the ground. The bones of mammals are well ossified and their teeth, which are usually differentiated, are coated in a layer of [[Tooth enamel|prismatic enamel]]. The teeth are shed once ([[milk teeth]]) during the animal's lifetime or not at all, as is the case in [[cetacea]]ns. Mammals have three bones in the middle ear and a [[cochlea]] in the [[inner ear]]. They are clothed in hair and their skin contains glands which secrete [[sweat gland|sweat]]. Some of these glands are specialized as [[mammary gland]]s, producing milk to feed the young. Mammals breathe with [[lung]]s and have a muscular [[Thoracic diaphragm|diaphragm]] separating the thorax from the abdomen which helps them draw air into the lungs. The mammalian heart has four chambers, and oxygenated and deoxygenated blood are kept entirely separate. Nitrogenous waste is excreted primarily as urea.<ref name=Dorit914>{{cite book |title=Zoology |url=https://archive.org/details/zoology0000dori/page/909 |url-access=registration |last1=Dorit |first1=R. L. |last2=Walker |first2=W. F. |last3=Barnes |first3=R. D. |year=1991 |publisher=Saunders College Publishing |isbn=978-0-03-030504-7 |pages=[https://archive.org/details/zoology0000dori/page/909 909–914] }}</ref> Mammals are [[amniote]]s, and most are [[viviparous]], giving birth to live young. Exceptions to this are the egg-laying [[monotreme]]s, the [[platypus]] and the [[echidna]]s of Australia. Most other mammals have a [[placenta]] through which the developing [[foetus]] obtains nourishment, but in [[marsupial]]s, the foetal stage is very short and the immature young is born and finds its way to its mother's [[Pouch (marsupial)|pouch]] where it latches on to a [[teat]] and completes its development.<ref name=Dorit914/> ====Human anatomy==== {{Further|Human body#Anatomy|Outline of human anatomy}} [[File:Parasagittal MRI of human head in patient with benign familial macrocephaly prior to brain injury (ANIMATED).gif|thumb|left|Sagittal sections of the head as seen by a modern [[MRI]] scan]] [[File:Violin Scroll (2769099541).jpg|upright|thumb|In humans, dexterous hand movements and increased brain size are likely to have evolved simultaneously.<ref name="BritBrit-Hand">{{cite encyclopedia |chapter-url=https://www.britannica.com/EBchecked/topic/254068/hand |title=Encyclopædia Britannica 2006 Ultimate Reference Suite DVD |chapter=Hand |access-date=15 May 2014 |archive-date=17 May 2014 |archive-url=https://web.archive.org/web/20140517132724/https://www.britannica.com/EBchecked/topic/254068/hand |url-status=live }}</ref>]] [[Human]]s have the overall body plan of a mammal. Humans have a head, neck, [[Trunk (anatomy)|trunk]] (which includes the [[thorax]] and [[abdomen]]), two arms and hands, and two legs and feet. Generally, students of certain [[biological sciences]], [[paramedic]]s, prosthetists and orthotists, [[physiotherapists]], [[occupational therapy|occupational therapists]], [[nurses]], [[podiatry|podiatrists]], and [[medical school|medical students]] learn gross anatomy and microscopic anatomy from anatomical models, skeletons, textbooks, diagrams, photographs, lectures and tutorials and in addition, medical students generally also learn gross anatomy through practical experience of [[dissection]] and inspection of [[cadaver]]s. The study of microscopic anatomy (or [[histology]]) can be aided by practical experience examining histological preparations (or slides) under a [[microscope]].<ref>{{cite web |url=https://www.medschoolsonline.co.uk/index.php?pageid=135 |title=Studying medicine |publisher=Medschools Online |access-date=27 June 2013 |archive-url=https://web.archive.org/web/20130128114829/https://www.medschoolsonline.co.uk/index.php?pageid=135 |archive-date=28 January 2013 |url-status=dead }}</ref> Human anatomy, physiology and biochemistry are complementary basic medical sciences, which are generally taught to medical students in their first year at medical school. Human anatomy can be taught regionally or systemically; that is, respectively, studying anatomy by bodily regions such as the head and chest, or studying by specific systems, such as the nervous or respiratory systems.<ref name="intro HGray" /> The major anatomy textbook, [[Gray's Anatomy]], has been reorganized from a systems format to a regional format, in line with modern teaching methods.<ref>{{cite book |url=https://archive.org/details/graysanatomyanat0000unse |title=Publisher's page for Gray's Anatomy. 39th edition (UK).|year=2004 |isbn=978-0-443-07168-3|url-access=registration |last1=Drake |first1=Richard Lee |last2=Gray |first2=Henry |last3=Vogl |first3=Wayne |last4=Mitchell |first4=Adam W. M. }}</ref><ref>{{cite book |url=https://archive.org/details/graysanatomyanat0000unse |title=Publisher's page for Gray's Anatomy. 39th edition (US).|year=2004 |isbn=978-0-443-07168-3|url-access=registration |last1=Drake |first1=Richard Lee |last2=Gray |first2=Henry |last3=Vogl |first3=Wayne |last4=Mitchell |first4=Adam W. M. }}</ref> A thorough working knowledge of anatomy is required by physicians, especially [[surgery|surgeons]] and doctors working in some diagnostic specialties, such as [[histopathology]] and [[radiology]].<ref name=AAA>{{cite web |url=https://www.anatomy.org/ |title=American Association of Anatomists |access-date=27 June 2013 |archive-url=https://web.archive.org/web/20190404042736/https://www.anatomy.org/ |archive-date=4 April 2019 |url-status=dead }}</ref> Academic anatomists are usually employed by universities, medical schools or teaching hospitals. They are often involved in teaching anatomy, and research into certain systems, organs, tissues or cells.<ref name=AAA/> == Invertebrate anatomy == [[File:Chirocephalus diaphanus male head.png|thumb|Head of a male ''[[Daphnia]]'', a [[planktonic]] crustacean]] [[Invertebrate]]s constitute a vast array of living organisms ranging from the simplest unicellular [[eukaryote]]s such as ''[[Paramecium]]'' to such complex multicellular animals as the [[octopus]], [[lobster]] and [[dragonfly]]. They constitute about 95% of the animal species. By definition, none of these creatures has a backbone. The cells of single-cell [[protozoa]]ns have the same basic structure as those of multicellular animals but some parts are specialized into the equivalent of tissues and organs. Locomotion is often provided by [[cilia]] or [[flagella]] or may proceed via the advance of [[pseudopodia]], food may be gathered by [[phagocytosis]], energy needs may be supplied by [[photosynthesis]] and the cell may be supported by an [[endoskeleton]] or an [[exoskeleton]]. Some protozoans can form multicellular colonies.<ref>{{cite book |title=Invertebrate Zoology, 7th edition |last1=Ruppert |first1=Edward E. |last2=Fox |first2=Richard, S. |last3=Barnes |first3=Robert D. |year=2004 |publisher=Cengage Learning |isbn=978-81-315-0104-7 |pages=23–24 }}</ref> [[Metazoa]]ns are a multicellular organism, with different groups of cells serving different functions. The most basic types of metazoan tissues are epithelium and connective tissue, both of which are present in nearly all invertebrates. The outer surface of the epidermis is normally formed of epithelial cells and secretes an [[extracellular matrix]] which provides support to the organism. An endoskeleton derived from the [[mesoderm]] is present in [[echinoderm]]s, [[sponge]]s and some [[cephalopod]]s. [[Exoskeleton]]s are derived from the epidermis and is composed of [[chitin]] in [[arthropod]]s (insects, spiders, ticks, shrimps, crabs, lobsters). [[Calcium carbonate]] constitutes the shells of [[molluscs]], [[brachiopod]]s and some tube-building [[polychaete worms]] and [[silica]] forms the exoskeleton of the microscopic [[diatom]]s and [[radiolaria]].<ref>{{cite encyclopedia |url=https://www.britannica.com/EBchecked/topic/198292/exoskeleton |title=Exoskeleton |encyclopedia=Encyclopædia Britannica |access-date=2 July 2013 |archive-date=3 May 2015 |archive-url=https://web.archive.org/web/20150503190050/https://www.britannica.com/EBchecked/topic/198292/exoskeleton |url-status=live }}</ref> Other invertebrates may have no rigid structures but the epidermis may secrete a variety of surface coatings such as the [[pinacoderm]] of sponges, the gelatinous cuticle of cnidarians ([[polyp (zoology)|polyp]]s, [[sea anemone]]s, [[jellyfish]]) and the [[collagen]]ous cuticle of [[annelid]]s. The outer epithelial layer may include cells of several types including sensory cells, gland cells and stinging cells. There may also be protrusions such as [[microvilli]], cilia, bristles, [[Spine (zoology)|spines]] and [[tubercle]]s.<ref>{{cite encyclopedia |url=https://www.britannica.com/EBchecked/topic/289723/integument |title=Integument |author=Ebling, F. J. G. |encyclopedia=Encyclopædia Britannica |access-date=2 July 2013 |archive-date=30 April 2015 |archive-url=https://web.archive.org/web/20150430013330/https://www.britannica.com/EBchecked/topic/289723/integument |url-status=live }}</ref> [[Marcello Malpighi]], the father of microscopical anatomy, discovered that plants had tubules similar to those he saw in insects like the silk worm. He observed that when a ring-like portion of bark was removed on a trunk a swelling occurred in the tissues above the ring, and he unmistakably interpreted this as growth stimulated by food coming down from the leaves, and being captured above the ring.<ref name="ArberAgnes">{{cite journal|author=Arber, Agnes|title= Nehemiah Grew (1641–1712) and Marcello Malpighi (1628–1694): an essay in comparison|journal= Isis |volume =34|issue=1| pages =7–16|jstor=225992|doi=10.1086/347742|year= 1942|s2cid= 143008947}}</ref> === Arthropod anatomy === {{main|Arthropod|Insect morphology|Spider anatomy}} [[Arthropod]]s comprise the largest phylum of [[invertebrate]]s in the animal kingdom with over a million known species.<ref>Britannica Concise Encyclopaedia 2007</ref> [[Insect]]s possess [[segmentation (biology)|segmented]] bodies supported by a hard-jointed outer covering, the [[exoskeleton]], made mostly of [[chitin]]. The segments of the body are organized into three distinct parts, a head, a [[Thorax (insect anatomy)|thorax]] and an [[abdomen]].<ref>{{cite web|title=O. Orkin Insect zoo |url=https://insectzoo.msstate.edu/Students/basic.structure.html |year=1997 |publisher=Mississippi State University |access-date=23 June 2013 |url-status=dead |archive-url=https://web.archive.org/web/20090602045832/https://www.insectzoo.msstate.edu/Students/basic.structure.html |archive-date=2 June 2009 }}</ref> The head typically bears a pair of sensory [[Antenna (biology)|antennae]], a pair of [[compound eye]]s, one to three simple eyes ([[ocelli]]) and three sets of modified appendages that form the [[insect mouthparts|mouthparts]]. The thorax has three pairs of segmented [[arthropod leg|legs]], one pair each for the three segments that compose the thorax and one or two pairs of [[insect wing|wings]]. The abdomen is composed of eleven segments, some of which may be fused and houses the [[digestion|digestive]], [[Respiration (physiology)|respiratory]], [[excretory]] and reproductive systems.<ref name="Gullan and Cranston">{{cite book |last1=Gullan |first1=P.J. |last2=Cranston |first2=P. S. |title=The Insects: An Outline of Entomology |publisher=Blackwell Publishing |location=Oxford |year=2005 |edition=3 |pages=[https://archive.org/details/isbn_9781405111133/page/22 22–48] |isbn=978-1-4051-1113-3 |url-access=registration |url=https://archive.org/details/isbn_9781405111133/page/22 }}</ref> There is considerable variation between species and many adaptations to the body parts, especially wings, legs, antennae and mouthparts.<ref>{{cite book |title=Invertebrate Zoology, 7th edition |last1=Ruppert |first1=Edward E. |last2=Fox |first2=Richard, S. |last3=Barnes |first3=Robert D. |year=2004 |publisher=Cengage Learning |isbn=978-81-315-0104-7 |pages=218–225 }}</ref> [[Spider]]s a class of [[arachnid]]s have four pairs of legs; a body of two segments—a [[cephalothorax]] and an [[abdomen]]. Spiders have no wings and no antennae. They have mouthparts called [[chelicerae]] which are often connected to venom glands as most spiders are venomous. They have a second pair of appendages called [[pedipalp]]s attached to the cephalothorax. These have similar segmentation to the legs and function as taste and smell organs. At the end of each male pedipalp is a spoon-shaped cymbium that acts to support the [[palpal bulb|copulatory organ]]. == Other branches of anatomy == * [[Surface anatomy]] is important as the study of anatomical landmarks that can be readily seen from the exterior contours of the body.<ref name="intro HGray"/> It enables medics and [[veterinarian]]s to gauge the position and anatomy of the associated deeper structures. Superficial is a directional term that indicates that structures are located relatively close to the surface of the body.<ref>{{cite book |last=Marieb |first=Elaine |title=Human Anatomy & Physiology |year=2010 |publisher=Pearson |location=San Francisco |page=12}}</ref> * [[Comparative anatomy]] relates to the comparison of anatomical structures (both gross and microscopic) in different animals.<ref name="intro HGray" /> * [[Artistic anatomy]] relates to anatomic studies of [[body proportions]] for artistic reasons. == History == {{main|History of anatomy}} === Ancient === [[File:Foetal positions in uterus, pregnant female Wellcome L0000845.jpg|thumb|upright=1.05|Image of early rendition of anatomy findings]] In 1600 BCE, the [[Edwin Smith Papyrus]], an [[Ancient Egyptian medicine|Ancient Egyptian]] [[Medical manual|medical text]], described the [[heart]] and its vessels, as well as the brain and its [[meninges]] and [[cerebrospinal fluid]], and the [[liver]], [[spleen]], [[kidneys]], [[uterus]] and [[bladder]]. It showed the [[blood vessel]]s diverging from the heart.<ref>{{Cite book |last=Rose |first=F. Clifford |chapter-url=https://books.google.com/books?id=2fkRDAAAQBAJ&pg=PA19 |title=Neurology and Trauma |date=2006-03-16 |publisher=Oxford University Press, USA |isbn=978-0-19-517032-0 |editor-last=Evans |editor-first=Randolph W. |language=en |chapter=The History of Cerebral Trauma |access-date=14 March 2023 |archive-date=26 March 2023 |archive-url=https://web.archive.org/web/20230326164813/https://books.google.com/books?id=2fkRDAAAQBAJ&pg=PA19 |url-status=live }}</ref><ref>{{Cite journal |last=Atta |first=Hussein M. |date=December 1999 |title=Edwin Smith Surgical Papyrus: The Oldest Known Surgical Treatise |url=http://journals.sagepub.com/doi/10.1177/000313489906501222 |journal=The American Surgeon |language=en |volume=65 |issue=12 |pages=1190–1192 |doi=10.1177/000313489906501222 |pmid=10597074 |s2cid=30179363 |issn=0003-1348 |access-date=7 March 2023 |archive-date=7 March 2023 |archive-url=https://web.archive.org/web/20230307133230/https://journals.sagepub.com/doi/10.1177/000313489906501222 |url-status=live }}</ref><ref>{{Cite journal |last1=Boehm |first1=Thomas |last2=Bleul |first2=Conrad C. |date=February 2007 |title=The evolutionary history of lymphoid organs |url=http://www.nature.com/articles/ni1435 |journal=Nature Immunology |language=en |volume=8 |issue=2 |pages=131–135 |doi=10.1038/ni1435 |pmid=17242686 |s2cid=45581056 |issn=1529-2908 |quote=Important landmark discoveries included the first description of the spleen found in the Edwin Smith Papyrus, containing medical information from Egypt dating back as early as 3000 BC... |access-date=7 March 2023 |archive-date=7 March 2023 |archive-url=https://web.archive.org/web/20230307140612/https://www.nature.com/articles/ni1435 |url-status=live }}</ref> The [[Ebers Papyrus]] ({{circa|1550 BCE}}) features a "treatise on the heart", with vessels carrying all the body's fluids to or from every member of the body.<ref>{{cite book | title = The Greatest Benefit to Mankind: A Medical History of Humanity from Antiquity to the Present | last = Porter | first = R. | year = 1997 | publisher = Harper Collins | isbn = 978-0-00-215173-3 | pages=49–50}}</ref> Ancient Greek anatomy and physiology underwent great changes and advances throughout the early medieval world. Over time, this medical practice expanded due to a continually developing understanding of the functions of organs and structures in the body. Phenomenal anatomical observations of the human body were made, which contributed to the understanding of the brain, eye, liver, reproductive organs, and nervous system. The [[Hellenistic Egypt]]ian city of [[Alexandria]] was the stepping-stone for Greek anatomy and physiology. Alexandria not only housed the biggest library for medical records and books of the liberal arts in the world during the time of the Greeks but was also home to many medical practitioners and philosophers. Great patronage of the arts and sciences from the [[Ptolemaic dynasty]] of Egypt helped raise Alexandria up, further rivalling other Greek states' cultural and scientific achievements.<ref name=Longrigg>{{cite journal|last1=Longrigg|first1=James|title=Anatomy in Alexandria in the Third Century B.C|journal=The British Journal for the History of Science|date=December 1988|volume=21|issue=4|pages=455–488|jstor=4026964|doi=10.1017/s000708740002536x|pmid=11621690|s2cid=37575399}}</ref> [[File:The Blue Beryl-Anatomy.jpg|thumb|An anatomy [[thangka]], part of [[Desi Sangye Gyatso]]'s The Blue Beryl, 17th century]] Some of the most striking advances in early anatomy and physiology took place in Hellenistic Alexandria.<ref name=Longrigg /> Two of the most famous anatomists and physiologists of the third century were [[Herophilus]] and [[Erasistratus]]. These two physicians helped pioneer human [[dissection]] for medical research, using the cadavers of condemned criminals, which was considered taboo until the Renaissance—Herophilus was recognized as the first person to perform systematic dissections.<ref name = bay>{{cite journal |last1= Bay|first1=Noel Si Yang|last2=Bay|first2=Boon-Huat|title=Greek Anatomists Herophilus: The Father of Anatomy|journal=Anatomy and Cell Biology|date=2010|volume= 43 |issue= 3|pages=280–283|doi=10.5115/acb.2010.43.4.280|pmc=3026179|pmid=21267401}}</ref> Herophilus became known for his anatomical works, making impressive contributions to many branches of anatomy and many other aspects of medicine.<ref>{{cite journal|last1=Von Staden|first1=H|title=The Discovery of the Body: Human Dissection and Its Cultural Contexts in Ancient Greece|journal=The Yale Journal of Biology and Medicine|date=1992|volume=65|issue=3|pages=223–241|pmid=1285450|pmc=2589595}}</ref> Some of the works included classifying the system of the pulse, the discovery that human arteries had thicker walls than veins, and that the atria were parts of the heart. Herophilus's knowledge of the human body has provided vital input towards understanding the brain, eye, liver, reproductive organs, and nervous system and characterizing the course of the disease.<ref name = bay/> Erasistratus accurately described the structure of the brain, including the cavities and membranes, and made a distinction between its cerebrum and cerebellum<ref>{{Cite web |url=https://www.faqs.org/health/bios/12/Erasistratus.html |title= Erasistratus Biography (304B.C-250B.C) |access-date=23 February 2022 |archive-date= 16 November 2018 |archive-url=https://web.archive.org/web/20181116054307/https://www.faqs.org/health/bios/12/Erasistratus.html |url-status=bot: unknown |website=Free Health Encyclopedia - faqs.org }}</ref> During his study in Alexandria, Erasistratus was particularly concerned with studies of the circulatory and nervous systems. He could distinguish the human body's sensory and motor nerves and believed air entered the lungs and heart, which was then carried throughout the body. His distinction between the arteries and veins—the arteries carrying the air through the body, while the veins carry the blood from the heart was a great anatomical discovery. Erasistratus was also responsible for naming and describing the function of the epiglottis and the heart's valves, including the tricuspid.<ref>{{cite encyclopedia|title=Erasistratus of Ceos: Greek Physician|date = April 3, 2018 |url= https://www.britannica.com/biography/Erasistratus-of-Ceos|encyclopedia= Encyclopædia Britannica|access-date=|archive-date=21 April 2019|archive-url= https://web.archive.org/web/20190421000007/https://www.britannica.com/biography/Erasistratus-of-Ceos |url-status=live}}</ref> During the third century, Greek physicians were able to differentiate nerves from blood vessels and tendons<ref>{{cite journal |last1= Wiltse|first1=LL|last2=Pait|first2=TG|title=Herophilus of Alexandria (325-255 B.C.) The Father of Anatomy|journal=Spine|date=1 September 1998|volume= 23 |issue= 17 |pages= 1904–1914|pmid=9762750|doi=10.1097/00007632-199809010-00022}}</ref> and to realize that the nerves convey neural impulses.<ref name="Longrigg"/> It was Herophilus who made the point that damage to motor nerves induced paralysis.<ref name = bay/> Herophilus named the meninges and ventricles in the brain, appreciated the division between cerebellum and cerebrum and recognized that the brain was the "seat of intellect" and not a "cooling chamber" as propounded by Aristotle<ref>{{cite journal|last1=Wills|first1=Adrian|title=Herophilus, Ersasistratus, and the birth of neuroscience|journal=The Lancet |date=1999 |volume= 354|issue=9191|pages=1719–1720|doi=10.1016/S0140-6736(99)02081-4|pmid=10568587|s2cid=30110082|url=https://www.thelancet.com/journals/lancet/article/PIIS0140-6736(99)02081-4/references |access-date=25 November 2015|archive-date=28 October 2019|archive-url=https://web.archive.org/web/20191028095251/https://www.thelancet.com/journals/lancet/article/PIIS0140-6736(99)02081-4/references |url-status=live}}</ref> Herophilus is also credited with describing the optic, oculomotor, motor division of the trigeminal, facial, vestibulocochlear and hypoglossal nerves.<ref name="Cambridge University Press">{{cite book|last1=Von Staden|first1=Heinrich |title=Herophilus: The Art of Medicine in Early Alexandria|date=October 2007|publisher=Cambridge University Press|isbn=9780521041782|url=https://www.cambridge.org/us/academic/subjects/classical-studies/ancient-philosophy/herophilus-art-medicine-early-alexandria-edition-translation-and-essays|access-date=25 November 2015|archive-date=8 December 2015|archive-url=https://web.archive.org/web/20151208053822/https://www.cambridge.org/us/academic/subjects/classical-studies/ancient-philosophy/herophilus-art-medicine-early-alexandria-edition-translation-and-essays|url-status=live}}</ref> [[File:Zahrawi1.png|200px|thumb|right|Surgical instruments were invented by [[Abulcasis]] in the 11th century]] [[File:Cheshm manuscript.jpg|200px|thumb|left|Anatomy of the eye for the first time in history by [[Hunayn ibn Ishaq]] in the 9th century]] [[File:13th century anatomical illustration.jpg|thumb|13th century anatomical illustration]] Incredible feats were made during the third century BCE in both the digestive and reproductive systems. Herophilus discovered and described not only the salivary glands but also the small intestine and liver.<ref name="Cambridge University Press"/> He showed that the uterus is a hollow organ and described the ovaries and uterine tubes. He recognized that spermatozoa were produced by the testes and was the first to identify the prostate gland.<ref name="Cambridge University Press"/> The anatomy of the muscles and skeleton is described in the ''[[Hippocratic Corpus]]'', an Ancient Greek medical work written by unknown authors.<ref>{{cite book |last=Gillispie |first=Charles Coulston | author-link=Charles Coulston Gillispie |title=Dictionary of Scientific Biography | volume=VI | pages=419–427 |year=1972 | publisher=Charles Scribner's Sons | location=New York}}</ref> [[Aristotle]] described [[vertebrate]] anatomy based on animal [[dissection]]. [[Praxagoras]] identified the difference between [[arteries]] and [[vein]]s. Also in the 4th century BCE, [[Herophilos]] and [[Erasistratus]] produced more accurate anatomical descriptions based on [[vivisection]] of criminals in [[Alexandria]] during the [[Ptolemaic Kingdom|Ptolemaic period]].<ref name=Bodies>{{cite book | url=https://books.google.com/books?id=5Xb9e3SLAZwC&q=Ptolemaic+vivisection&pg=PA256 | title=Medicine and Society in Ptolemaic Egypt | author=Lang, Philippa | publisher=Brill NV | year=2013 | page=256 | isbn=978-9004218581 | access-date=15 October 2020 | archive-date=16 April 2021 | archive-url=https://web.archive.org/web/20210416082236/https://books.google.com/books?id=5Xb9e3SLAZwC&q=Ptolemaic+vivisection&pg=PA256 | url-status=live }}</ref><ref>[https://exhibits.hsl.virginia.edu/antiqua/alexandrian/ "Alexandrian Medicine"] {{Webarchive|url=https://web.archive.org/web/20170220135427/https://exhibits.hsl.virginia.edu/antiqua/alexandrian/ |date=20 February 2017 }}. ''Antiqua Medicina – from Homer to Vesalius''. University of Virginia.</ref> In the 2nd century, [[Galen of Pergamum]], an anatomist, [[clinician]], writer, and philosopher,<ref name="BritBrit-Galen"/> wrote the final and highly influential anatomy treatise of ancient times.<ref name="pmid1081972">{{cite journal |vauthors=Charon NW, Johnson RC, Muschel LH | title = Antileptospiral activity in lower-vertebrate sera | journal = Infect. Immun. | volume = 12 | issue = 6 | pages = 1386–1391 | year = 1975 | pmid = 1081972 | pmc = 415446 | doi = 10.1128/IAI.12.6.1386-1391.1975}}</ref> He compiled existing knowledge and studied anatomy through the dissection of animals.<ref name="BritBrit-Galen">{{cite encyclopedia |chapter-url=https://www.britannica.com/EBchecked/topic/223895/Galen-of-Pergamum |title=Encyclopædia Britannica 2006 Ultimate Reference Suite DVD |chapter=Galen of Pergamum |first=Vivien |last=Hutton |access-date=13 May 2014 |archive-date=6 April 2012 |archive-url=https://web.archive.org/web/20120406224329/https://www.britannica.com/EBchecked/topic/223895/Galen-of-Pergamum |url-status=live }}</ref> He was one of the first experimental physiologists through his [[vivisection]] experiments on animals.<ref>Brock, Arthur John (translator) ''[https://archive.org/stream/galenonnaturalfa00galeuoft#page/xxxii/mode/2up Galen. On the Natural Faculties]''. Edinburgh, 1916. Introduction, page xxxiii.</ref> Galen's drawings, based mostly on dog anatomy, became effectively the only anatomical textbook for the next thousand years.<ref name=Boas/> His work was known to [[Renaissance]] doctors only through [[Islamic Golden Age]] medicine until it was translated from Greek sometime in the 15th century.<ref name=Boas/> === Medieval to early modern === [[File:Leonardo da Vinci - RCIN 919000, Verso The bones and muscles of the arm c.1510-11.jpg|left|thumb|Anatomical study of the arm, by [[Leonardo da Vinci]], (about 1510)]] [[File:Charta ex qva figvram parare convenit, illi qvae nervorvm seriem exprimit appendendam, 1543..JPG|thumb|upright|Anatomical chart in [[Vesalius]]'s ''Epitome'', 1543]] [[File:Michiel Jansz van Mierevelt - Anatomy lesson of Dr. Willem van der Meer.jpg|thumb|right|[[Michiel Jansz van Mierevelt]] – ''Anatomy lesson of Dr. Willem van der Meer'', 1617]] Anatomy developed little from classical times until the sixteenth century; as the historian Marie Boas writes, "Progress in anatomy before the sixteenth century is as mysteriously slow as its development after 1500 is startlingly rapid".<ref name=Boas>{{cite book | title=The Scientific Renaissance 1450–1630 | publisher=Fontana | author=Boas, Marie | year=1970 |orig-year=first published by Collins, 1962 | pages=120–143}}</ref>{{rp|120–121}} Between 1275 and 1326, the anatomists [[Mondino de Luzzi]], [[Alessandro Achillini]] and [[Antonio Benivieni]] at [[Bologna]] carried out the first systematic human dissections since ancient times.<ref name="ZimmermanVeith1993">{{cite book | last1=Zimmerman | first1=Leo M. | last2=Veith | first2=Ilza | title=Great Ideas in the History of Surgery | url=https://books.google.com/books?id=ABbCI7z4UwMC | year=1993 | publisher=Norman | isbn=978-0-930405-53-3 | access-date=31 July 2017 | archive-date=15 April 2016 | archive-url=https://web.archive.org/web/20160415082135/https://books.google.com/books?id=ABbCI7z4UwMC | url-status=live }}</ref><ref name="Crombie1959">{{cite book | last=Crombie | first=Alistair Cameron | title=The History of Science From Augustine to Galileo | url=https://books.google.com/books?id=bGDScHy1clsC&pg=PA4 | year=1959 | publisher=Courier Dover Publications | isbn=978-0-486-28850-5 | access-date=31 July 2017 | archive-date=9 April 2016 | archive-url=https://web.archive.org/web/20160409055609/https://books.google.com/books?id=bGDScHy1clsC&pg=PA4 | url-status=live }}</ref><ref name="Thorndike1958">{{cite book | last=Thorndike | first=Lynn | title=A History of Magic and Experimental Science: Fourteenth and fifteenth centuries | url=https://books.google.com/books?id=IbvlQFj4YfUC&pg=PA586 | year=1958 | publisher=Columbia University Press | isbn=978-0-231-08797-1 | access-date=31 July 2017 | archive-date=16 April 2016 | archive-url=https://web.archive.org/web/20160416061340/https://books.google.com/books?id=IbvlQFj4YfUC&pg=PA586 | url-status=live }}</ref> Mondino's ''Anatomy'' of 1316 was the first textbook in the medieval rediscovery of human anatomy. It describes the body in the order followed in Mondino's dissections, starting with the abdomen, thorax, head, and limbs. It was the standard anatomy textbook for the next century.<ref name=Boas/> [[Leonardo da Vinci]] (1452–1519) was trained in anatomy by [[Andrea del Verrocchio]].<ref name=Boas/> He made use of his anatomical knowledge in his artwork, making many sketches of skeletal structures, muscles and organs of humans and other vertebrates that he dissected.<ref name=Boas/><ref>{{cite book | last=Mason | first=Stephen F. | title=A History of the Sciences | url=https://archive.org/details/historyofscience00maso | url-access=registration | publisher=Collier | year=1962 | location = New York | page=[https://archive.org/details/historyofscience00maso/page/550 550]}}</ref> [[Andreas Vesalius]] (1514–1564), professor of anatomy at the [[University of Padua]], is considered the founder of modern human anatomy.<ref>{{cite web | url=https://www2.warwick.ac.uk/newsandevents/pressreleases/new_material_from/ | title=Warwick honorary professor explores new material from founder of modern human anatomy | work=Press release | publisher=University of Warwick | access-date=8 July 2013 | archive-date=6 November 2018 | archive-url=https://web.archive.org/web/20181106231146/https://warwick.ac.uk/newsandevents/pressreleases/new_material_from/ | url-status=live }}</ref> Originally from [[Duchy of Brabant|Brabant]], Vesalius published the influential book ''[[De humani corporis fabrica]]'' ("the structure of the human body"), a large format book in seven volumes, in 1543.<ref>Vesalius, Andreas. ''De humani corporis fabrica libri septem''. Basileae [Basel]: ''Ex officina'' Joannis Oporini, 1543.</ref> The accurate and intricately detailed illustrations, often in [[allegorical]] poses against Italianate landscapes, are thought to have been made by the artist [[Jan van Calcar]], a pupil of [[Titian]].<ref>O'Malley, C.D. ''Andreas Vesalius of Brussels, 1514–1564''. Berkeley: University of California Press, 1964.</ref> In England, anatomy was the subject of the first public lectures given in any science; these were provided by the [[Barber surgeon|Company of Barbers and Surgeons]] in the 16th century, joined in 1583 by the Lumleian lectures in surgery at the [[Royal College of Physicians]].<ref name=Boas229>{{cite book | title=The Scientific Renaissance 1450–1630 | publisher=Fontana | author=Boas, Marie | year=1970 |orig-year=first published by Collins, 1962 | page=229}}</ref> === Late modern === {{further|History of anatomy in the 19th century}} [[File:Lektion_i_anatomi_vid_Gymnastiska_Centralinstitutet_Stockholm_kvinnliga_kursen_1891-1893_gih0124.jpg|left|thumb|Anatomy teaching with female students, 1891–1893]] Medical schools began to be set up in the United States towards the end of the 18th century. Classes in anatomy needed a continual stream of cadavers for dissection, and these were difficult to obtain. Philadelphia, Baltimore, and New York were all renowned for [[body snatching]] activity as criminals raided graveyards at night, removing newly buried corpses from their coffins.<ref name=trafficdead>{{cite book |author=Sappol, Michael |title=A traffic of dead bodies: anatomy and embodied social identity in nineteenth-century America |publisher=Princeton University Press |location=Princeton, NJ |year=2002 |isbn=978-0-691-05925-9 |url=https://books.google.com/books?id=-9cKRzEx6ywC&q=A+Traffic+of+Dead+Bodies |access-date=15 October 2020 |archive-date=16 April 2021 |archive-url=https://web.archive.org/web/20210416090946/https://books.google.com/books?id=-9cKRzEx6ywC&q=A+Traffic+of+Dead+Bodies |url-status=live }}</ref> A similar problem existed in Britain where demand for bodies became so great that grave-raiding and even [[anatomy murder]] were practised to obtain cadavers.<ref name="Rosner, Lisa. 2010">Rosner, Lisa. 2010. The Anatomy Murders. Being the True and Spectacular History of Edinburgh's Notorious Burke and Hare and of the Man of Science Who Abetted Them in the Commission of Their Most Heinous Crimes. University of Pennsylvania Press</ref> Some graveyards were, in consequence, protected with watchtowers. The practice was halted in Britain by the [[Anatomy Act]] of 1832,<ref>{{cite book | title=Death, Dissection, and the Destitute | publisher=Penguin | author=Richardson, Ruth | year=1989 | isbn=978-0-14-022862-5}}</ref><ref>{{cite web | url=https://www.leeds.ac.uk/chb/lectures/anatomy1.html | title=Introductory Anatomy | publisher=University of Leeds | access-date=25 June 2013 | author=Johnson, D.R. | archive-date=4 November 2008 | archive-url=https://web.archive.org/web/20081104162600/https://www.leeds.ac.uk/chb/lectures/anatomy1.html | url-status=live }}</ref> while in the United States, similar legislation was enacted after the physician [[William S. Forbes]] of [[Jefferson Medical College]] was found guilty in 1882 of "complicity with resurrectionists in the despoliation of graves in Lebanon Cemetery".<ref>{{cite web|url=https://www.jefferson.edu/about/eakins/forbes.html |title=Reproduction of Portrait of Professor William S. Forbes |publisher=Jefferson: Eakins Gallery |access-date=14 October 2013 |url-status=dead |archive-url=https://web.archive.org/web/20131016064638/https://www.jefferson.edu/about/eakins/forbes.html |archive-date=16 October 2013}}</ref> The teaching of anatomy in Britain was transformed by Sir [[John Struthers (anatomist)|John Struthers]], [[Regius Professor of Anatomy (Aberdeen)|Regius Professor of Anatomy]] at the [[University of Aberdeen]] from 1863 to 1889. He was responsible for setting up the system of three years of "pre-clinical" academic teaching in the sciences underlying medicine, including especially anatomy. This system lasted until the reform of medical training in 1993 and 2003. As well as teaching, he collected many vertebrate skeletons for his museum of [[comparative anatomy]], published over 70 research papers, and became famous for his public dissection of the [[Tay Whale]].<ref name=pmid17373426>{{cite journal |vauthors=Waterston SW, Laing MR, Hutchison JD | title = Nineteenth century medical education for tomorrow's doctors | journal = Scottish Medical Journal | volume = 52 | issue = 1 | pages = 45–49 | year = 2007 | pmid = 17373426 | doi=10.1258/rsmsmj.52.1.45| s2cid = 30286930 }}</ref><ref name="pmid15712576">{{cite journal |vauthors=Waterston SW, Hutchison JD | title = Sir John Struthers MD FRCS Edin LLD Glasg: Anatomist, zoologist and pioneer in medical education | journal = The Surgeon | volume = 2 | issue = 6 | pages = 347–351 | year = 2004 | pmid = 15712576 | doi=10.1016/s1479-666x(04)80035-0}}</ref> From 1822 the Royal College of Surgeons regulated the teaching of anatomy in medical schools.<ref name="McLachlan, J. 2006. p.243-53">{{cite journal | author = McLachlan J., Patten D. | year = 2006 | title = Anatomy teaching: ghosts of the past, present and future | journal = Medical Education | volume = 40 | issue = 3| pages = 243–253 | doi = 10.1111/j.1365-2929.2006.02401.x | pmid = 16483327 | s2cid = 30909540 }}</ref> Medical museums provided examples in comparative anatomy, and were often used in teaching.<ref>{{cite journal | author = Reinarz J | year = 2005 | title = The age of museum medicine: The rise and fall of the medical museum at Birmingham's School of Medicine | journal = Social History of Medicine | volume = 18 | issue = 3| pages = 419–437 | doi = 10.1093/shm/hki050 }}</ref> [[Ignaz Semmelweis]] investigated [[puerperal fever]] and he discovered how it was caused. He noticed that the frequently fatal fever occurred more often in mothers examined by medical students than by midwives. The students went from the dissecting room to the hospital ward and examined women in childbirth. Semmelweis showed that when the trainees washed their hands in chlorinated lime before each clinical examination, the incidence of puerperal fever among the mothers could be reduced dramatically.<ref>{{cite encyclopedia |url=https://global.britannica.com/EBchecked/topic/534198/Ignaz-Philipp-Semmelweis |title=Ignaz Philipp Semmelweis |encyclopedia=Encyclopædia Britannica |access-date=15 October 2013}}</ref> [[File:Siemens-electron-microscope.jpg|upright|thumb|An electron microscope from 1973]] Before the modern medical era, the primary means for studying the internal structures of the body were [[dissection]] of the dead and [[inspection]], [[palpation]], and [[auscultation]] of the living. The advent of [[microscopy]] opened up an understanding of the building blocks that constituted living tissues. Technical advances in the development of [[achromatic lens]]es increased the [[Angular resolution|resolving power]] of the microscope, and around 1839, [[Matthias Jakob Schleiden]] and [[Theodor Schwann]] identified that cells were the fundamental unit of organization of all living things. The study of small structures involved passing light through them, and the [[microtome]] was invented to provide sufficiently thin slices of tissue to examine. Staining techniques using artificial dyes were established to help distinguish between different tissue types. Advances in the fields of [[histology]] and [[cytology]] began in the late 19th century<ref name=BritMicro>{{cite encyclopedia |url=https://www.britannica.com/EBchecked/topic/22980/anatomy/283/Microscopic-anatomy |title=Microscopic anatomy |encyclopedia=Encyclopædia Britannica |access-date=14 October 2013 |archive-date=28 October 2014 |archive-url=https://web.archive.org/web/20141028075812/https://www.britannica.com/EBchecked/topic/22980/anatomy/283/Microscopic-anatomy |url-status=live }}</ref> along with advances in surgical techniques allowing for the painless and safe removal of [[biopsy]] specimens. The invention of the [[electron microscope]] brought a significant advance in resolution power and allowed research into the [[ultrastructure]] of cells and the [[organelle]]s and other structures within them. About the same time, in the 1950s, the use of [[X-ray diffraction]] for studying the crystal structures of proteins, nucleic acids, and other biological molecules gave rise to a new field of [[molecular anatomy]].<ref name=BritMicro/> Equally important advances have occurred in ''non-invasive'' techniques for examining the body's interior structures. [[X-ray]]s can be passed through the body and used in medical [[radiography]] and [[fluoroscopy]] to differentiate interior structures that have varying degrees of opaqueness. [[Magnetic resonance imaging]], [[computed tomography]], and [[ultrasound imaging]] have all enabled the examination of internal structures in unprecedented detail to a degree far beyond the imagination of earlier generations.<ref name=":1">{{cite web | url=https://www.mhhe.com/biosci/ap/foxhumphys/student/olc/h-reading1.html | title=Anatomical Imaging | publisher=McGraw Hill Higher Education | year=1998 | access-date=25 June 2013 | archive-url=https://web.archive.org/web/20160303232044/https://www.mhhe.com/biosci/ap/foxhumphys/student/olc/h-reading1.html | archive-date=3 March 2016 | url-status=dead }}</ref> == See also == * [[Anatomical model]] * {{section link|Bibliography of biology|Anatomy}} * [[Outline of human anatomy]] * [[Plastination]] * [[Evelyn tables]] * {{Portal inline|size=tiny|Anatomy}} == References == {{reflist}} == External links == {{Library resources box}} * [https://www.bbc.co.uk/podcasts/series/iots/all#playepisode115 Anatomy], ''[[In Our Time (BBC Radio 4)|In Our Time]]''. [[BBC Radio 4]]. [[Melvyn Bragg]] with guests Ruth Richardson, Andrew Cunningham and [[Harold Ellis (professor)|Harold Ellis]]. * [https://www.bartleby.com/107/ "Anatomy of the Human Body". 20th edition. 1918. Henry Gray] * {{cite EB1911 |wstitle=Anatomy |volume=1 |pages=920–943 |short=1 |first=Frederick Gymer |last=Parsons }} * [https://anatomia.library.utoronto.ca Anatomia Collection: anatomical plates 1522 to 1867] (digitized books and images) *Lyman, Henry Munson. ''[https://digital.sciencehistory.org/works/bg257g476 The Book of Health]'' (1898). [https://digital.sciencehistory.org/ Science History Institute Digital Collections] {{Webarchive|url=https://web.archive.org/web/20190202042542/https://digital.sciencehistory.org/ |date=2 February 2019 }}. * Gunther von Hagens ''[https://vonhagens-plastination.com/pages/medical-teaching-specimens/von-hagens-plastination.php/silicone-plastinates True Anatomy for New Ways of Teaching]''. == Sources == {{Free-content attribution| | title = Openstax Anatomy and Physiology | author = J. 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