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countermeasures</span> </div> </a> <button aria-controls="toc-Constraints_and_countermeasures-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Constraints and countermeasures subsection</span> </button> <ul id="toc-Constraints_and_countermeasures-sublist" class="vector-toc-list"> <li id="toc-Environmental_conditions_and_social_interactions_affecting_microbial_cheating" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Environmental_conditions_and_social_interactions_affecting_microbial_cheating"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>Environmental conditions and social interactions affecting microbial cheating</span> </div> </a> <ul id="toc-Environmental_conditions_and_social_interactions_affecting_microbial_cheating-sublist" class="vector-toc-list"> </ul> </li> <li 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.mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">For other uses, see <a href="/wiki/Cheating_(disambiguation)" class="mw-disambig" title="Cheating (disambiguation)">cheating (disambiguation)</a>.</div> <p><b>Cheating</b> is a term used in <a href="/wiki/Behavioral_ecology" title="Behavioral ecology">behavioral ecology</a> and <a href="/wiki/Ethology" title="Ethology">ethology</a> to describe behavior whereby organisms receive a benefit at the cost of other organisms. Cheating is common in many mutualistic and altruistic relationships.<sup id="cite_ref-Bronstein2002_1-0" class="reference"><a href="#cite_note-Bronstein2002-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> A cheater is an individual who does not cooperate (or cooperates less than their fair share) but can potentially gain the benefit from others cooperating.<sup id="cite_ref-West2006_2-0" class="reference"><a href="#cite_note-West2006-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Cheaters are also those who selfishly use common resources to maximize their individual fitness at the expense of a group.<sup id="cite_ref-MacLean2006_3-0" class="reference"><a href="#cite_note-MacLean2006-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a> favors cheating, but there are mechanisms to regulate it.<sup id="cite_ref-Kokko_4-0" class="reference"><a href="#cite_note-Kokko-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> The <a href="/wiki/Stress_gradient_hypothesis" title="Stress gradient hypothesis">stress gradient hypothesis</a> states that facilitation, cooperation or mutualism should be more common in stressful environments, while cheating, competition or parasitism are common in benign environments (i.e nutrient excess). </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Theoretical_models">Theoretical models</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cheating_(biology)&action=edit&section=1" title="Edit section: Theoretical models"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Organisms communicate and cooperate to perform a wide range of behaviors. <a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">Mutualism</a>, or mutually beneficial interactions between species, is common in ecological systems.<sup id="cite_ref-Bronstein_5-0" class="reference"><a href="#cite_note-Bronstein-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> These interactions can be thought of "biological markets" in which species offer partners goods that are relatively inexpensive for them to produce and receive goods that are more expensive or even impossible for them to produce.<sup id="cite_ref-Schwartz1998_6-0" class="reference"><a href="#cite_note-Schwartz1998-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> However, these systems provide opportunities for exploitation by individuals that can obtain resources while providing nothing in return. Exploiters can take on several forms: individuals outside a mutualistic relationship who obtain a commodity in a way that confers no benefit to either mutualist, individuals who receive benefits from a partner but have lost the ability to give any in return, or individuals who have the option of behaving mutualistically towards their partners but chose not to do so.<sup id="cite_ref-Bronstein_5-1" class="reference"><a href="#cite_note-Bronstein-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>Cheaters, who do not cooperate but benefit from others who do cooperate gain a competitive edge. In an evolutionary context, this competitive edge refers to a greater ability to survive or to reproduce. If individuals who cheat are able to gain survivorship and reproductive benefits while incurring no costs, natural selection should favor cheaters. However, mechanisms also exist that prevent cheaters from undermining mutualistic systems. One main factor is that the advantages of cheating are often frequency-dependent. <a href="/wiki/Frequency-dependent_selection" title="Frequency-dependent selection">Frequency-dependent selection</a> occurs when the <a href="/wiki/Fitness_(biology)" title="Fitness (biology)">fitness</a> of a phenotype depends on its frequency relative to other phenotypes in a population. Cheater phenotypes often display negative frequency-dependent selection, where fitness increases as a phenotype becomes less common and vice versa.<sup id="cite_ref-Riechert2009_7-0" class="reference"><a href="#cite_note-Riechert2009-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> In other words, cheaters do best (in terms of evolutionary benefits such as increased survival and reproduction) when there are relatively few of them, but as cheaters become more abundant, they do worse. </p><p>For example, in <i>Escherichia coli</i> colonies, there are antibiotic-sensitive "cheaters" that persist at low numbers on antibiotic-laced mediums when in a cooperative colony. These cheaters enjoy the benefit of others producing antibiotic-resistant agents while producing none themselves. However, as numbers increase, if they persist in not producing the antibiotic agent themselves, they are more likely to be negatively impacted by the antibiotic substrate because there is less antibiotic agent to protect everyone.<sup id="cite_ref-Riechert2009_7-1" class="reference"><a href="#cite_note-Riechert2009-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> Thus, cheaters can persist in a population because their exploitative behavior gives them an advantage when they exist at low frequencies but these benefits are diminished when they are greater in number. </p><p>Others have proposed that cheating (exploitive behavior) can stabilize cooperation in mutualistic systems.<sup id="cite_ref-Kokko_4-1" class="reference"><a href="#cite_note-Kokko-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Little2009_8-0" class="reference"><a href="#cite_note-Little2009-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> In many mutualistic systems, there will be feedback benefits to those that cooperate. For instance, the fitness of both partners may be improved. If there is a high reward or many benefits for the individual that initiated the cooperative behavior, mutualism should be selected for. When researchers investigated the co-evolution of cooperation and choice in a choosy host and its symbiont (an organism that lives in a relationship that benefits all parties involved), their model indicated that although choice and cooperation may be initially selected for, this would often be unstable.<sup id="cite_ref-Kokko_4-2" class="reference"><a href="#cite_note-Kokko-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> In other words, one cooperative partner will choose another cooperative partner if given a choice. However, if this choice is made over and over, variation is removed and this selection can no longer be maintained. This situation is similar to the <a href="/wiki/Lek_mating" title="Lek mating">lek paradox</a> in female choice. For example, in lek paradox, if females consistently choose for a particular male trait, genetic variance for that trait should eventually be eliminated, removing the benefits of choice. However, that choice somehow still persists. </p><p>One theory states that cheating maintains genetic variation in the face of selection for mutualism. One study shows that a small influx of immigrants with a tendency to cooperate less can generate enough genetic variability to stabilize selection for mutualism.<sup id="cite_ref-Kokko_4-3" class="reference"><a href="#cite_note-Kokko-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> This suggests that the presence of exploitive individuals, otherwise known as cheaters, contribute enough genetic variation to maintain mutualism itself. Both this theory and the negative frequency-dependent theory suggest that that cheating exists as part of a stable mixed evolutionary strategy with mutualism. In other words, cheating is a stable strategy used by individuals in a population where many other individuals cooperate. Another study supports that cheating can exist as a mixed strategy with mutualism using a mathematical game model.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> Thus, cheating can arise and be maintained in mutualistic populations. </p> <div class="mw-heading mw-heading2"><h2 id="Examples">Examples</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cheating_(biology)&action=edit&section=2" title="Edit section: Examples"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Stalked_slime_mould_fruiting_bodies_-_geograph.org.uk_-_923299.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c3/Stalked_slime_mould_fruiting_bodies_-_geograph.org.uk_-_923299.jpg/220px-Stalked_slime_mould_fruiting_bodies_-_geograph.org.uk_-_923299.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c3/Stalked_slime_mould_fruiting_bodies_-_geograph.org.uk_-_923299.jpg/330px-Stalked_slime_mould_fruiting_bodies_-_geograph.org.uk_-_923299.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c3/Stalked_slime_mould_fruiting_bodies_-_geograph.org.uk_-_923299.jpg/440px-Stalked_slime_mould_fruiting_bodies_-_geograph.org.uk_-_923299.jpg 2x" data-file-width="640" data-file-height="480" /></a><figcaption>Stalked slime mould fruiting bodies</figcaption></figure> <p>Studies of cheating and dishonest communication in populations presupposes an organismal system that cooperates. Without a collective population that has signaling and interactions among individuals, behaviors such as cheating do not manifest. In other words, in order to study cheating behavior, a model system that engages in <a href="/wiki/Cooperation" title="Cooperation">cooperation</a> is needed. Models that provide insight on cheating include the social amoeba <i><a href="/wiki/Dictyostelium_discoideum" title="Dictyostelium discoideum">Dictyostelium discoideum</a></i>;<sup id="cite_ref-Strassmann2000_10-0" class="reference"><a href="#cite_note-Strassmann2000-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Santorelli2008_11-0" class="reference"><a href="#cite_note-Santorelli2008-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Li2011_12-0" class="reference"><a href="#cite_note-Li2011-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Eusociality" title="Eusociality">eusocial</a> insects, such as ants, bees, and wasps;<sup id="cite_ref-Wenseleers2004a_13-0" class="reference"><a href="#cite_note-Wenseleers2004a-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> and inter-specific interactions found in cleaning mutualisms. Common examples of cleaning mutualisms include <a href="/wiki/Cleaner_fish" title="Cleaner fish">cleaner fish</a> such as wrasses and gobies,<sup id="cite_ref-Soares2014_14-0" class="reference"><a href="#cite_note-Soares2014-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bshary2002_15-0" class="reference"><a href="#cite_note-Bshary2002-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> and some <a href="/wiki/Cleaner_shrimp" title="Cleaner shrimp">cleaner shrimp</a>.<sup id="cite_ref-Chapuis2009_16-0" class="reference"><a href="#cite_note-Chapuis2009-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="In_Dictyostelium_discoideum">In <i>Dictyostelium discoideum</i></h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cheating_(biology)&action=edit&section=3" title="Edit section: In Dictyostelium discoideum"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><i><a href="/wiki/Dictyostelium_discoideum" title="Dictyostelium discoideum">Dictyostelium discoideum</a></i> is a widely used model for cooperation and the development of <a href="/wiki/Multicellular_organism" title="Multicellular organism">multicellularity</a>. This species of amoeba is most commonly found in a <a href="/wiki/Ploidy" title="Ploidy">haploid</a>, single-celled state that feeds independently and undergoes <a href="/wiki/Asexual_reproduction" title="Asexual reproduction">asexual reproduction</a>. However, when the scarcity of food sources causes individual cells to starve, roughly 10⁴ to 10⁵ cells aggregate to form a mobile, multicellular structure dubbed a "slug".<sup id="cite_ref-Strassmann2000_10-1" class="reference"><a href="#cite_note-Strassmann2000-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> In the wild, aggregates generally contain multiple genotypes, resulting in chimeric mixtures. Unlike clonal (genetically identical) aggregates typically found in multicellular organisms, the potential for competition exists in chimeric aggregates.<sup id="cite_ref-Strassmann2000_10-2" class="reference"><a href="#cite_note-Strassmann2000-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Santorelli2008_11-1" class="reference"><a href="#cite_note-Santorelli2008-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Li2011_12-1" class="reference"><a href="#cite_note-Li2011-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> For example, because individuals in the aggregate contain different genomes, differences in fitness can result in conflict of interest among cells in the aggregate, where different genotypes could potentially compete against each other for resources and reproduction.<sup id="cite_ref-Strassmann2000_10-3" class="reference"><a href="#cite_note-Strassmann2000-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> In <i>Dictyostelium discoideum</i>, roughly 20% of the cells in the aggregate become dead to make the stalk of a <a href="/wiki/Sorocarp" title="Sorocarp">fruiting body</a>. The remaining 80% of cells become spores in the sorus of the fruiting body, which can germinate again once conditions are more favorable.<sup id="cite_ref-Strassmann2000_10-4" class="reference"><a href="#cite_note-Strassmann2000-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Santorelli2008_11-2" class="reference"><a href="#cite_note-Santorelli2008-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> In this case, 20% of the cells must give up reproduction so that the fruiting body forms successfully. This makes chimeric aggregates of <i>Dictyostelium discoideum</i> susceptible to cheating individuals that take advantage of the reproductive behavior without paying the fair price. In other words, if certain individuals tend to become a part of the sorus more frequently, they can gain increased benefit from the fruiting body system without sacrificing their own opportunities to reproduce.<sup id="cite_ref-Strassmann2000_10-5" class="reference"><a href="#cite_note-Strassmann2000-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Santorelli2008_11-3" class="reference"><a href="#cite_note-Santorelli2008-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Li2011_12-2" class="reference"><a href="#cite_note-Li2011-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> Cheating behavior in <i>D. discoideum</i> is well established, and many studies have attempted to elucidate the evolutionary and genetic mechanisms underlying the behavior. Having a 34Mb genome that is completely sequenced and well annotated makes <i>D. discoideum</i> a useful model in studying the genetic bases and molecular mechanisms of cheating, and in a broader sense, <a href="/wiki/Evolution_of_eusociality" title="Evolution of eusociality">social evolution</a>.<sup id="cite_ref-Li2011_12-3" class="reference"><a href="#cite_note-Li2011-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="In_eusocial_insects">In eusocial insects</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cheating_(biology)&action=edit&section=4" title="Edit section: In eusocial insects"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Eusocial insects also serve as valuable tools in studying cheating. Eusocial insects behave cooperatively, where members of the community forgo reproduction to assist a few individuals to reproduce. Such model systems have potential for conflict of interest to arise among individuals, and thus also have potential for cheating to occur.<sup id="cite_ref-Wenseleers2004a_13-1" class="reference"><a href="#cite_note-Wenseleers2004a-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Cheney2005_17-0" class="reference"><a href="#cite_note-Cheney2005-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> Eusocial insects in the order Hymenoptera, which includes bees and wasps, exhibit good examples of conflicts of interest present in insect societies. In these systems, queen bees and wasps can mate and lay fertilized eggs that hatch into females. On the other hand, workers of most species in Hymenoptera can produce eggs, but cannot produce fertilized eggs due to loss of mating ability.<sup id="cite_ref-Wenseleers2004a_13-2" class="reference"><a href="#cite_note-Wenseleers2004a-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> Workers that lay eggs represent a cost for the colony because workers that lay eggs often do significantly less work, and thus negatively impact the health of the colony (for example: decreased amount of collected food, or less attention given to tending the queen's eggs). In this case, a conflict of interest arises between the workers and the colony. The workers should lay eggs in order to pass on their genes; however, as a colony, having only the queen reproduce leads to better productivity.<sup id="cite_ref-Wenseleers2004a_13-3" class="reference"><a href="#cite_note-Wenseleers2004a-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> If workers sought to pass their own genes by laying eggs, foraging activities would diminish, leading to decreased resources for the entire colony. This, in turn, can cause a <a href="/wiki/Tragedy_of_the_commons" title="Tragedy of the commons">tragedy of the commons</a>,<sup id="cite_ref-Wenseleers2004b_18-0" class="reference"><a href="#cite_note-Wenseleers2004b-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> where selfish behavior lead to the depletion of resources, with long-term negative consequences for the group. However, in natural bee and wasp societies, only 0.01–0.1% and 1%, respectively, of the workers lay eggs, suggesting that strategies exist to combat cheating to prevent tragedy of the commons.<sup id="cite_ref-Wenseleers2004a_13-4" class="reference"><a href="#cite_note-Wenseleers2004a-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Wenseleers2004b_18-1" class="reference"><a href="#cite_note-Wenseleers2004b-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> These insect systems have given scientists opportunities to study strategies that keep cheating in check. Such strategies are commonly referred to as "policing" strategies, generally where additional costs are imposed on cheaters to discourage or eliminate cheating behaviors. For example, honeybees and wasps may eat eggs produced by workers. In some ant species and yellowjackets, policing may occur via aggression towards or killing egg-laying individuals to minimize cheating.<sup id="cite_ref-Wenseleers2004a_13-5" class="reference"><a href="#cite_note-Wenseleers2004a-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Cleaning_station_konan.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/98/Cleaning_station_konan.jpg/220px-Cleaning_station_konan.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/98/Cleaning_station_konan.jpg/330px-Cleaning_station_konan.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/98/Cleaning_station_konan.jpg/440px-Cleaning_station_konan.jpg 2x" data-file-width="1777" data-file-height="1333" /></a><figcaption>Fish cleaned by smaller cleaner wrasses on Hawaiian reefs</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="In_cleaning_symbiosis">In cleaning symbiosis</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cheating_(biology)&action=edit&section=5" title="Edit section: In cleaning symbiosis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Cleaning_symbiosis" title="Cleaning symbiosis">Cleaning symbiosis</a> that develop between small and larger marine organisms often represent models useful for studying the evolution of stable social interactions and cheating. In the cleaning fish <i>Labroides dimidiatus</i> (<a href="/wiki/Bluestreak_cleaner_wrasse" title="Bluestreak cleaner wrasse">Bluestreak cleaner wrasse</a>), as in many cleaner species, client fish seeks to have <a href="/wiki/Parasitism" title="Parasitism">ectoparasites</a> removed by the cleaners. In these situations, instead of picking off the parasites on the surface of the client fish, the cleaner can cheat by feeding on the client's tissue (mucus layer, scales, etc.), thereby gaining additional benefit from the symbiotic system.<sup id="cite_ref-Bshary2002_15-1" class="reference"><a href="#cite_note-Bshary2002-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> It has been well documented that cleaners will feed on mucus when their clients are unable to control the cleaner's behavior; however, in natural settings, client fish often jolt, chase after cheating cleaners, or terminate interactions of swimming way, effectively controlling the cheating behavior.<sup id="cite_ref-Bshary2002_15-2" class="reference"><a href="#cite_note-Bshary2002-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Chapuis2009_16-1" class="reference"><a href="#cite_note-Chapuis2009-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Cheney2005_17-1" class="reference"><a href="#cite_note-Cheney2005-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> Studies on cleaning mutualisms generally suggest that cheating behavior is often adjusted depending on the species of the client. In cleaning shrimp, cheating is predicted to occur less often because shrimps bear a higher cost if the clients use aggression to control the cleaner's behavior.<sup id="cite_ref-Chapuis2009_16-2" class="reference"><a href="#cite_note-Chapuis2009-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> Studies have found that cleaner species can strategically adjust cheating behavior according to the potential associated risk. For example, predatory clients, which present a significantly high cost for cheating, experience less cheating behavior. On the other hand, nonpredatory clients present a lower cost for cheating, and thus experience more cheating behaviors from the cleaners.<sup id="cite_ref-Chapuis2009_16-3" class="reference"><a href="#cite_note-Chapuis2009-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Cheney2005_17-2" class="reference"><a href="#cite_note-Cheney2005-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> Some evidence suggest that physiological processes can mediate the cleaners' decision to switch from cooperating to cheating in mutualistic interactions. For example, in the bluestreak cleaner wrasse, changes in <a href="/wiki/Cortisol" title="Cortisol">cortisol</a> levels are associated with behavior changes.<sup id="cite_ref-Soares2014_14-1" class="reference"><a href="#cite_note-Soares2014-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> For smaller clients, increase cortisol levels in the water lead to more cooperative behavior, while for larger clients, the same treatment lead to more dishonest behavior. It has been suggested that "good behavior" toward smaller clients often allow wrasses to attract larger clients that are often cheated.<sup id="cite_ref-Soares2014_14-2" class="reference"><a href="#cite_note-Soares2014-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Other">Other</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cheating_(biology)&action=edit&section=6" title="Edit section: Other"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Other models of cheating include the <a href="/wiki/European_tree_frog" title="European tree frog">European tree frog</a>, "Hyla arborea". In many sexually reproducing species such as this, some males can access mates by exploiting resources of more competitive males.<sup id="cite_ref-Brepson2012_19-0" class="reference"><a href="#cite_note-Brepson2012-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> Many species have dynamic reproductive strategies that can change in response to changes in the environment. In these instances, several factors contribute to the decision to switch between mating strategies. For example, in the European tree frog, a sexually competitive (as in, perceived to be attractive by females) male tend to call to attract mates. This is often referred to as the "bourgeois" tactic.<sup id="cite_ref-Brepson2012_19-1" class="reference"><a href="#cite_note-Brepson2012-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> On the other hand, a smaller male that would likely fail to attract mates using the bourgeois tactic will tend to hide near attractive males and attempt to access females. In this instance, the males can gain access to females without having to defend territories or acquiring additional resources (which often serve as the basis for attractiveness). This is referred to as the "parasitic" tactic, where the smaller male effectively cheats its way to accessing females, by reaping the benefit of sexual reproduction without contributing resources that normally attract females.<sup id="cite_ref-Brepson2012_19-2" class="reference"><a href="#cite_note-Brepson2012-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> Models such as this provide valuable tools for research aimed at energetic constraints and <a href="/wiki/Sensory_cue" title="Sensory cue">environmental cues</a> involved in cheating. Studies find that mating strategies are highly adaptable and depend on a variety of factors, such as competitiveness, energetic costs involved in defending territory or acquiring resources.<sup id="cite_ref-Brepson2012_19-3" class="reference"><a href="#cite_note-Brepson2012-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Constraints_and_countermeasures">Constraints and countermeasures</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cheating_(biology)&action=edit&section=7" title="Edit section: Constraints and countermeasures"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Environmental_conditions_and_social_interactions_affecting_microbial_cheating">Environmental conditions and social interactions affecting microbial cheating</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cheating_(biology)&action=edit&section=8" title="Edit section: Environmental conditions and social interactions affecting microbial cheating"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Like many other organisms, bacteria rely on iron intake for its biological processes.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="removed citation to predatory publisher content (December 2019)">citation needed</span></a></i>]</sup> However, iron is sometimes difficult to access in certain environments, like soil. Some bacteria have evolved <a href="/wiki/Siderophore" title="Siderophore">siderophores</a>, iron-chelating particles that seek and bring back iron for the bacteria. Siderophores are not necessarily specific to its producer - sometimes another individual could take up the particles instead.<sup id="cite_ref-:0_22-0" class="reference"><a href="#cite_note-:0-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> <i><a href="/wiki/Pseudomonas_fluorescens" title="Pseudomonas fluorescens">Pseudomonas fluorescens</a></i> is a bacterium commonly found in the soil. Under low-iron conditions, <i>P. fluorescens</i> produces siderophores, specifically <a href="/wiki/Pyoverdine" title="Pyoverdine">pyoverdine</a>, to retrieve the iron necessary for survival. However, when iron is readily available, either from freely diffusing in environment or another bacterium's siderophores, <i>P. fluorescens</i> ceases production, allowing the bacterium to devote its energy towards growth. One study showed that when <i>P. fluorescens</i> grew in association with <i><a href="/wiki/Streptomyces_ambofaciens" title="Streptomyces ambofaciens">Streptomyces ambofaciens</a></i>, another bacterium that produces the siderophore <a href="/w/index.php?title=Coelichelin&action=edit&redlink=1" class="new" title="Coelichelin (page does not exist)">coelichelin</a>, no pyoverdine was detected. This result suggested that <i>P. fluorescens</i> ceased siderophore production in favor of taking up iron-bound coelichelin, an association also known as siderophore piracy.<sup id="cite_ref-:0_22-1" class="reference"><a href="#cite_note-:0-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p><p>More studies, however, suggested that <i>P. fluorescens'</i> cheating behavior could be suppressed. In another study, two strains of <i>P. fluorescens</i> were studied in the soil, their natural environment. One strain, known as the producer, produced a higher level of siderophores, which meant that other strain, known as the non-producer, ceased siderophore production in favor of using the other's siderophores. Although one would expect that the non-producer would outcompete the producer, like the <i>P. fluorescens</i> and <i>S. ambofaciens</i> association, the study demonstrated that the non-producer was unable to do so in soil conditions, suggesting that the two strains could coexist. Further experiments suggested that this cheating prevention may be due to interactions with other microbes in the soil influencing the relationship or the spatial structure of the soil preventing siderophore diffusion and therefore limiting the non-producer's ability to exploit the producer's siderophores.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Selection_pressure_in_bacteria_(intraspecies)"><span id="Selection_pressure_in_bacteria_.28intraspecies.29"></span>Selection pressure in bacteria (intraspecies)</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cheating_(biology)&action=edit&section=9" title="Edit section: Selection pressure in bacteria (intraspecies)"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>By definition, individuals cheat to gain benefits that their non-cheating counterparts do not receive. This raises the question of how a cooperative system can exist in face of these cheaters. One answer is that the cheaters actually have a reduced fitness compared to the non-cheaters. </p><p>In a study by Dandekar et al., the researchers examined the survival rates of cheating and non-cheating bacteria populations (<i><a href="/wiki/Pseudomonas_aeruginosa" title="Pseudomonas aeruginosa">Pseudomonas aeruginosa</a></i>) under varying environmental conditions.<sup id="cite_ref-Dandekar2012_24-0" class="reference"><a href="#cite_note-Dandekar2012-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> These microorganisms, like many species of bacteria, use a cell-cell communication system called <a href="/wiki/Quorum_sensing" title="Quorum sensing">quorum sensing</a> that detect their population density and prompt the transcription of various resources when needed. In this case, the resources are publicly shared proteases that break down a food source like casein, and privately used adenosine hydrolase, which breaks down another food source, adenosine. The problem arises when some individuals ("cheaters") do not respond to these quorum sensing signals and therefore do not contribute to the costly protease production yet enjoy the benefits of the broken down resources. </p><p>When <i>P. aeruginosa</i> populations are placed into growth conditions where cooperation (and responding to the quorum signal) is costly, the number of cheaters increases, and the public resources are depleted, which can lead to a <a href="/wiki/Tragedy_of_the_commons" title="Tragedy of the commons">tragedy of the commons</a>. However, when <i>P. aeruginosa</i> populations are placed into growth conditions with a proportion of adenosine, the cheaters are suppressed because the bacteria that responds to the quorum signal now produces adenosine hydrolase that they privately use for themselves to digest adenosine food source. In wild populations where the presence of adenosine is common, this is an explanation for how individuals that cooperate could have higher fitness than those that cheat, thereby suppressing the cheaters and maintaining cooperation. </p> <div class="mw-heading mw-heading3"><h3 id="Policing/punishment_in_insects"><span id="Policing.2Fpunishment_in_insects"></span>Policing/punishment in insects</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cheating_(biology)&action=edit&section=10" title="Edit section: Policing/punishment in insects"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Worker_policing" title="Worker policing">Worker policing</a></div> <p>Cheating is also commonly found in insects. The social and seemingly altruistic communities found in insects such as ants and bees provide ample opportunities for cheaters to take advantage of the system and accrue additional benefits at the expense of the community. </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Wasp_nest_3.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0e/Wasp_nest_3.jpg/220px-Wasp_nest_3.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0e/Wasp_nest_3.jpg/330px-Wasp_nest_3.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0e/Wasp_nest_3.jpg/440px-Wasp_nest_3.jpg 2x" data-file-width="3456" data-file-height="2592" /></a><figcaption>Wasp nest, with some larvae</figcaption></figure> <p>Sometimes, a colony of insects is called a "<a href="/wiki/Superorganism" title="Superorganism">superorganism</a>" for its ability to take on properties greater than those of the sum of individuals. A colony of insects in which different individuals are specialized for specific tasks means a greater colony production and greater efficiency.<sup id="cite_ref-Ratnieks2006_25-0" class="reference"><a href="#cite_note-Ratnieks2006-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> Moreover, based on the <a href="/wiki/Kin_selection" title="Kin selection">kin-selection theory</a>, it is collectively beneficial for all the individuals in the community to have the queen to lay eggs rather than the workers lay eggs.<sup id="cite_ref-Beekman2008_26-0" class="reference"><a href="#cite_note-Beekman2008-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> This is because if the workers lay eggs, it benefits the egg-laying worker individually, but the rest of the workers are now twice removed from this worker's offspring. Therefore, though it is beneficial for one individual to have its own offspring, it is collectively beneficial to have the queen lay the eggs. Therefore, a system of worker and queen policing exists against worker-laid eggs. </p><p>One form of policing occurs by the oophagy of the worker-laid eggs, found in many ant and bee species.<sup id="cite_ref-Ratnieks2006_25-1" class="reference"><a href="#cite_note-Ratnieks2006-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> This could be done by both or either the queen or the workers. In a series of experiments with honeybees (<i><a href="/wiki/Western_honey_bee" title="Western honey bee">Apis mellifera</a></i>), Ratneiks & Visscher found that other workers effectively removed worker-laid eggs in all colonies, whether the eggs were from originated from the same colonies or not.<sup id="cite_ref-Ratnieks1989_27-0" class="reference"><a href="#cite_note-Ratnieks1989-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> An example of a combination of queen and worker policing is found in ants, in the genus <i><a href="/wiki/Diacamma" title="Diacamma">Diacamma</a></i>, in which worker-laid eggs are taken by other workers and fed to the "queen".<sup id="cite_ref-Kikuta1999_28-0" class="reference"><a href="#cite_note-Kikuta1999-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> In general, these signals that identify the eggs as queen-laid are likely incorruptible, since it must be an honest signal to be maintained and not be used by cheating workers.<sup id="cite_ref-Beekman2008_26-1" class="reference"><a href="#cite_note-Beekman2008-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p><p>The other form of policing occurs through aggression towards egg-laying workers. In a species of tree wasp <i><a href="/wiki/Dolichovespula_sylvestris" title="Dolichovespula sylvestris">Dolichovespula sylvestris</a></i>, Wenseleers et al. found that a combination of aggressive behavior and destruction of worker-laid eggs kept the number of worker-laid eggs low.<sup id="cite_ref-Wenseleers2005_29-0" class="reference"><a href="#cite_note-Wenseleers2005-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> In fact, 91% of the worker-laid eggs were policed within one day. They also found that about 20% of workers laying eggs were prevented from doing so through both the queen's and workers' aggressive behavior. The workers and the queen would grab the egg-laying worker and try to sting her or push her off the cell. This usually results in the worker removing her abdomen and not depositing her eggs. </p> <div class="mw-heading mw-heading3"><h3 id="Policing/punishment_in_other_organisms"><span id="Policing.2Fpunishment_in_other_organisms"></span>Policing/punishment in other organisms</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cheating_(biology)&action=edit&section=11" title="Edit section: Policing/punishment in other organisms"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Nest_of_naked_mole_rats.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Nest_of_naked_mole_rats.jpg/220px-Nest_of_naked_mole_rats.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Nest_of_naked_mole_rats.jpg/330px-Nest_of_naked_mole_rats.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Nest_of_naked_mole_rats.jpg/440px-Nest_of_naked_mole_rats.jpg 2x" data-file-width="2816" data-file-height="2112" /></a><figcaption>A nest of naked mole rats</figcaption></figure> <p>Aggression and punishment are not just found in insects. For example, in <a href="/wiki/Naked_mole-rat" title="Naked mole-rat">naked mole rats</a>, punishments by the queen are a way she motivates the lazier, less-related workers in their groups.<sup id="cite_ref-Reeve1992_30-0" class="reference"><a href="#cite_note-Reeve1992-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> The queen would shove the lazier workers, with the number of shoves increasing when there are fewer active workers. Reeve found that if the queen is removed when colonies are satiated, there is a significant drop in weight of the active workers because the lazier workers are taking advantage of the system. </p><p>Punishment is also a method used by cichlid <i><a href="/w/index.php?title=Neolamprologous_pulcher&action=edit&redlink=1" class="new" title="Neolamprologous pulcher (page does not exist)">Neolamprologous pulcher</a></i> in their cooperative breeding systems. It is a pay-to-stay system where helper fish are allowed to stay in certain territories in exchange for their help.<sup id="cite_ref-Fischer2014_31-0" class="reference"><a href="#cite_note-Fischer2014-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> Similar to the naked mole rats, the helpers that were prevented from helping, the "idle helpers", receive more aggression than control helpers in the study. Researchers theorize that this system developed because the fish are usually not closely related (so kinship benefits have little impact), and because there is a high level of predation risk when the fish is outside the group (therefore a strong motivator for the helper fish to stay in the group).<sup id="cite_ref-Fischer2014_31-1" class="reference"><a href="#cite_note-Fischer2014-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Rhesus_macaque" title="Rhesus macaque">Rhesus monkeys</a> also use aggression as a punishment. These animals have five distinct calls that they can "decide" to produce upon finding food. Whether they call or not is related to their <a href="/wiki/Sex" title="Sex">sex</a> and number of kin: females call more often and females with more kin call more often. However, sometimes when food is found, the individual ("discoverer") does not call to attract its kin and, presumably, to share food. If lower ranked individuals find this discoverer to be in the food drop area of the experiment, they recruit coalition support against this individual by screaming. The formed coalition then chases this individual away. If higher ranked individuals find this discoverer, they either chase the discoverer away or became physically aggressive towards the individual.<sup id="cite_ref-Hauser1992_32-0" class="reference"><a href="#cite_note-Hauser1992-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> These results show that aggression as punishment is a way to encourage members to work together and share food when it is found. </p> <div class="mw-heading mw-heading3"><h3 id="Interspecific_countermeasures">Interspecific countermeasures</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cheating_(biology)&action=edit&section=12" title="Edit section: Interspecific countermeasures"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Cheating and constraints of cheating are not limited to intraspecific interactions; it can also occur in a mutualistic relationship between two species. A common example is the mutualistic relationship between cleaner fish <i><a href="/wiki/Bluestreak_cleaner_wrasse" title="Bluestreak cleaner wrasse">Labroides dimidiatus</a></i> and reef fish. Bshary and Grutter found that cleaner wrasse prefers the client tissue mucus over ectoparasites.<sup id="cite_ref-Bshary2005_33-0" class="reference"><a href="#cite_note-Bshary2005-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> This creates a conflict between the cleaner fish and reef fish, because the reef fish only benefit when the cleaner fish eats the ectoparasites. Further studies revealed that in a lab setting, the cleaner fish undergoes behavioral change in face of deterrents against eating their preferential food.<sup id="cite_ref-Bshary2005_33-1" class="reference"><a href="#cite_note-Bshary2005-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> In several trials, the plate of their preferential food source was immediately removed when they eat it, to mimic "client fleeing" in natural settings. In other trials, the plate of their preferential food source chased the cleaner fish when they eat it, mimicking "client chasing" in natural setting. After only six learning trials, the cleaners learned to choose against their preference, indicating that punishment is potentially a very effective countermeasure against cheating in mutualistic relationships. </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Nitrogen-fixing_nodules_in_the_roots_of_legumes..JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/20/Nitrogen-fixing_nodules_in_the_roots_of_legumes..JPG/220px-Nitrogen-fixing_nodules_in_the_roots_of_legumes..JPG" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/20/Nitrogen-fixing_nodules_in_the_roots_of_legumes..JPG/330px-Nitrogen-fixing_nodules_in_the_roots_of_legumes..JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/20/Nitrogen-fixing_nodules_in_the_roots_of_legumes..JPG/440px-Nitrogen-fixing_nodules_in_the_roots_of_legumes..JPG 2x" data-file-width="2048" data-file-height="1365" /></a><figcaption>Nitrogen-fixing nodules in legumes</figcaption></figure> <p>Finally the countermeasures are not limited to organismal relationships. West et al. found a similar countermeasure against cheating in legume-rhizobium mutualism.<sup id="cite_ref-West2002_34-0" class="reference"><a href="#cite_note-West2002-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> In this relationship, nitrogen fixing bacteria <a href="/wiki/Rhizobia" title="Rhizobia">rhizobium</a> fixes atmospheric N<sub>2</sub> from inside the roots of leguminous plants, providing this essential source of nitrogen to these plants while also receiving organic acids for themselves. However, some bacteria are more mutualistic, while others are more parasitic because they consume the plant's resources but fixes little to no N<sub>2</sub>. Moreover, these plants cannot tell whether the bacteria are more or less parasitic until they are settled in the plant nodules. To prevent cheating, these plants seem to be able to punish the rhizobium bacteria. In a series of experiments, researchers forced non-cooperation between the bacteria and the plants by placing various nodules in nitrogen-free atmosphere. They saw a decrease in the rhizobium reproductive success by 50%.<sup id="cite_ref-Kiers2003_35-0" class="reference"><a href="#cite_note-Kiers2003-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> West et al. created a model for legume sanctioning the bacteria and hypothesizes that these behaviors exist to stabilize mutualistic interactions.<sup id="cite_ref-West2002_34-1" class="reference"><a href="#cite_note-West2002-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> </p><p>Another well-known example of plant-organism interaction occurs between <a href="/wiki/Yucca" title="Yucca">yuccas</a> and <a href="/wiki/Prodoxidae" title="Prodoxidae">yucca moths</a>. The female yucca moths deposit their eggs one at a time to the yucca flower. At the same time, she also deposits a small amount of pollen from yucca flowers as nutrition for the yucca moths. Because most of the pollen is not consumed by the larva, yucca moths are therefore also the active pollinators for the yucca plant. Moreover, sometimes the female moths do not successfully deposit their eggs the first time, and may try again and again. The yucca plant receives scars from the multiple attempts, but they also receive more pollen, since the moths are depositing pollen with every try. </p><p>"Cheating" sometimes happens when the yucca moth deposits too many eggs in one plant. In this case, the yucca plant has little to no benefits from this interaction. However, the plant has a unique way of constraining this behavior. While the constraint against cheating often occurs directly to the individual, in this case, the constraint occurs to the individual's offspring. The yucca plant can "abort" the moths by aborting the flowers. Pellmyr and Huth found that there is selective maturation for flowers that have low egg loads and high number of scars (and therefore a high amount of pollen).<sup id="cite_ref-Pellmyr1994_36-0" class="reference"><a href="#cite_note-Pellmyr1994-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> In this way, there is selection against the "cheaters" who try to use the yucca plant without providing the benefits of pollination. </p> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cheating_(biology)&action=edit&section=13" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-Bronstein2002-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-Bronstein2002_1-0">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFFerriere,_R.Bronstein,_J.L.Rinaldi,_S.Law,_R.2002" class="citation journal cs1">Ferriere, R.; Bronstein, J.L.; Rinaldi, S.; Law, R.; Gauduchon, M. (2002). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1690960">"Cheating and the evolutionary stability of mutualisms"</a>. <i>Proc. R. Soc. 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.navbar{float:left;text-align:left;margin-right:0.5em}body.skin--responsive .mw-parser-output .navbox-image img{max-width:none!important}@media print{body.ns-0 .mw-parser-output .navbox{display:none!important}}</style></div><div role="navigation" class="navbox" aria-labelledby="Evolutionary_biology" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><style data-mw-deduplicate="TemplateStyles:r1239400231">.mw-parser-output .navbar{display:inline;font-size:88%;font-weight:normal}.mw-parser-output .navbar-collapse{float:left;text-align:left}.mw-parser-output .navbar-boxtext{word-spacing:0}.mw-parser-output .navbar ul{display:inline-block;white-space:nowrap;line-height:inherit}.mw-parser-output .navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar li{word-spacing:-0.125em}.mw-parser-output .navbar a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}html.skin-theme-clientpref-night .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Evolution" title="Template:Evolution"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Evolution" title="Template talk:Evolution"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Evolution" title="Special:EditPage/Template:Evolution"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Evolutionary_biology" style="font-size:114%;margin:0 4em"><a href="/wiki/Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/wiki/Introduction_to_evolution" title="Introduction to evolution">Introduction</a></li> <li><a href="/wiki/Outline_of_evolution" title="Outline of evolution">Outline</a></li> <li><a href="/wiki/Timeline_of_the_evolutionary_history_of_life" title="Timeline of the evolutionary history of life">Timeline of evolution</a></li> <li><a href="/wiki/History_of_life" title="History of life">History of life</a></li> <li><a href="/wiki/Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolution" title="Evolution">Evolution</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Abiogenesis" title="Abiogenesis">Abiogenesis</a></li> <li><a href="/wiki/Adaptation" title="Adaptation">Adaptation</a></li> <li><a href="/wiki/Adaptive_radiation" title="Adaptive radiation">Adaptive radiation</a></li> <li><a href="/wiki/Altruism_(biology)" title="Altruism (biology)">Altruism</a> <ul><li><a class="mw-selflink selflink">Cheating</a></li> <li><a href="/wiki/Reciprocal_altruism" title="Reciprocal altruism">Reciprocal</a></li></ul></li> <li><a href="/wiki/Baldwin_effect" title="Baldwin effect">Baldwin effect</a></li> <li><a href="/wiki/Cladistics" title="Cladistics">Cladistics</a></li> <li><a href="/wiki/Coevolution" title="Coevolution">Coevolution</a> <ul><li><a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">Mutualism</a></li></ul></li> <li><a href="/wiki/Common_descent" title="Common descent">Common descent</a></li> <li><a href="/wiki/Convergent_evolution" title="Convergent evolution">Convergence</a></li> <li><a href="/wiki/Divergent_evolution" title="Divergent evolution">Divergence</a></li> <li><a href="/wiki/Earliest_known_life_forms" title="Earliest known life forms">Earliest known life forms</a></li> <li><a href="/wiki/Evidence_of_common_descent" title="Evidence of common descent">Evidence of evolution</a></li> <li><a href="/wiki/Evolutionary_arms_race" title="Evolutionary arms race">Evolutionary arms race</a></li> <li><a href="/wiki/Evolutionary_pressure" title="Evolutionary pressure">Evolutionary pressure</a></li> <li><a href="/wiki/Exaptation" title="Exaptation">Exaptation</a></li> <li><a href="/wiki/Extinction" title="Extinction">Extinction</a> <ul><li><a href="/wiki/Extinction_event" title="Extinction event">Event</a></li></ul></li> <li><a href="/wiki/Homology_(biology)" title="Homology (biology)">Homology</a></li> <li><a href="/wiki/Last_universal_common_ancestor" title="Last universal common ancestor">Last universal common ancestor</a></li> <li><a href="/wiki/Macroevolution" title="Macroevolution">Macroevolution</a></li> <li><a href="/wiki/Microevolution" title="Microevolution">Microevolution</a></li> <li><a href="/wiki/Evolutionary_mismatch" title="Evolutionary mismatch">Mismatch</a></li> <li><a href="/wiki/Nonadaptive_radiation" title="Nonadaptive radiation">Non-adaptive radiation</a></li> <li><a href="/wiki/Abiogenesis" title="Abiogenesis">Origin of life</a></li> <li><a href="/wiki/Panspermia" title="Panspermia">Panspermia</a></li> <li><a href="/wiki/Parallel_evolution" title="Parallel evolution">Parallel evolution</a></li> <li><a href="/wiki/Signalling_theory" title="Signalling theory">Signalling theory</a> <ul><li><a href="/wiki/Handicap_principle" title="Handicap principle">Handicap principle</a></li></ul></li> <li><a href="/wiki/Speciation" title="Speciation">Speciation</a> <ul><li><a href="/wiki/Species" title="Species">Species</a></li> <li><a href="/wiki/Species_complex" title="Species complex">Species complex</a></li></ul></li> <li><a href="/wiki/Evolutionary_taxonomy" title="Evolutionary taxonomy">Taxonomy</a></li> <li><a href="/wiki/Unit_of_selection" title="Unit of selection">Unit of selection</a> <ul><li><a href="/wiki/Gene-centered_view_of_evolution" title="Gene-centered view of evolution">Gene-centered view of evolution</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Population_genetics" title="Population genetics">Population<br />genetics</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Selective_breeding" title="Selective breeding">Artificial selection</a></li> <li><a href="/wiki/Biodiversity" title="Biodiversity">Biodiversity</a></li> <li><a href="/wiki/Evolutionarily_stable_strategy" title="Evolutionarily stable strategy">Evolutionarily stable strategy</a></li> <li><a href="/wiki/Fisher%27s_principle" title="Fisher's principle">Fisher's principle</a></li> <li><a href="/wiki/Fitness_(biology)" title="Fitness (biology)">Fitness</a> <ul><li><a href="/wiki/Inclusive_fitness" title="Inclusive fitness">Inclusive</a></li></ul></li> <li><a href="/wiki/Gene_flow" title="Gene flow">Gene flow</a></li> <li><a href="/wiki/Genetic_drift" title="Genetic drift">Genetic drift</a></li> <li><a href="/wiki/Kin_selection" title="Kin selection">Kin selection</a> <ul><li><a href="/wiki/Parental_investment" title="Parental investment">Parental investment</a></li> <li><a href="/wiki/Parent%E2%80%93offspring_conflict" title="Parent–offspring conflict">Parent–offspring conflict</a></li></ul></li> <li><a href="/wiki/Mutation" title="Mutation">Mutation</a></li> <li><a href="/wiki/Population" title="Population">Population</a></li> <li><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></li> <li><a href="/wiki/Sexual_dimorphism" title="Sexual dimorphism">Sexual dimorphism</a></li> <li><a href="/wiki/Sexual_selection" title="Sexual selection">Sexual selection</a> <ul><li><a href="/wiki/Sexual_selection_in_flowering_plants" title="Sexual selection in flowering plants">Flowering plants</a></li> <li><a href="/wiki/Sexual_selection_in_fungi" title="Sexual selection in fungi">Fungi</a></li> <li><a href="/wiki/Mate_choice" title="Mate choice">Mate choice</a></li></ul></li> <li><a href="/wiki/Social_selection" title="Social selection">Social selection</a></li> <li><a href="/wiki/Trivers%E2%80%93Willard_hypothesis" title="Trivers–Willard hypothesis">Trivers–Willard hypothesis</a></li> <li><a href="/wiki/Genetic_variation" title="Genetic variation">Variation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Development</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Canalisation_(genetics)" title="Canalisation (genetics)">Canalisation</a></li> <li><a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evolutionary developmental biology</a></li> <li><a href="/wiki/Genetic_assimilation" title="Genetic assimilation">Genetic assimilation</a></li> <li><a href="/wiki/Inversion_(evolutionary_biology)" title="Inversion (evolutionary biology)">Inversion</a></li> <li><a href="/wiki/Modularity_(biology)" title="Modularity (biology)">Modularity</a></li> <li><a href="/wiki/Phenotypic_plasticity" title="Phenotypic plasticity">Phenotypic plasticity</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Taxon" title="Taxon">taxa</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_bacteria" title="Evolution of bacteria">Bacteria</a></li> <li><a href="/wiki/Evolution_of_birds" title="Evolution of birds">Birds</a> <ul><li><a href="/wiki/Origin_of_birds" title="Origin of birds">origin</a></li></ul></li> <li><a href="/wiki/Evolution_of_brachiopods" title="Evolution of brachiopods">Brachiopods</a></li> <li><a href="/wiki/Evolution_of_molluscs" title="Evolution of molluscs">Molluscs</a> <ul><li><a href="/wiki/Evolution_of_cephalopods" title="Evolution of cephalopods">Cephalopods</a></li></ul></li> <li><a href="/wiki/Dinosaur#Evolutionary_history" title="Dinosaur">Dinosaurs</a></li> <li><a href="/wiki/Evolution_of_fish" title="Evolution of fish">Fish</a></li> <li><a href="/wiki/Evolution_of_fungi" title="Evolution of fungi">Fungi</a></li> <li><a href="/wiki/Evolution_of_insects" title="Evolution of insects">Insects</a> <ul><li><a href="/wiki/Evolution_of_butterflies" title="Evolution of butterflies">butterflies</a></li></ul></li> <li><a href="/wiki/History_of_life" title="History of life">Life</a></li> <li><a href="/wiki/Evolution_of_mammals" title="Evolution of mammals">Mammals</a> <ul><li><a href="/wiki/Cat_gap" title="Cat gap">cats</a></li> <li><a href="/wiki/Canidae#Evolution" title="Canidae">canids</a> <ul><li><a href="/wiki/Evolution_of_the_wolf" title="Evolution of the wolf">wolves</a></li> <li><a href="/wiki/Domestication_of_the_dog" title="Domestication of the dog">dogs</a></li></ul></li> <li><a href="/wiki/Hyena#Evolution" title="Hyena">hyenas</a></li> <li><a href="/wiki/Evolution_of_cetaceans" title="Evolution of cetaceans">dolphins and whales</a></li> <li><a href="/wiki/Evolution_of_the_horse" title="Evolution of the horse">horses</a></li> <li><a href="/wiki/Evolution_of_Macropodidae" title="Evolution of Macropodidae">Kangaroos</a></li> <li><a href="/wiki/Evolution_of_primates" title="Evolution of primates">primates</a> <ul><li><a href="/wiki/Human_evolution" title="Human evolution">humans</a></li> <li><a href="/wiki/Evolution_of_lemurs" title="Evolution of lemurs">lemurs</a></li></ul></li> <li><a href="/wiki/Evolution_of_sirenians" title="Evolution of sirenians">sea cows</a></li></ul></li> <li><a href="/wiki/Evolutionary_history_of_plants" title="Evolutionary history of plants">Plants</a> <ul><li><a href="/wiki/Pollinator-mediated_selection" title="Pollinator-mediated selection">pollinator-mediated</a></li></ul></li> <li><a href="/wiki/Evolution_of_reptiles" title="Evolution of reptiles">Reptiles</a></li> <li><a href="/wiki/Evolution_of_spiders" title="Evolution of spiders">Spiders</a></li> <li><a href="/wiki/Evolution_of_tetrapods" title="Evolution of tetrapods">Tetrapods</a></li> <li><a href="/wiki/Viral_evolution" title="Viral evolution">Viruses</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Organ_(anatomy)" class="mw-redirect" title="Organ (anatomy)">organs</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_cells" title="Evolution of cells">Cell</a></li> <li><a href="/wiki/Models_of_DNA_evolution" title="Models of DNA evolution">DNA</a></li> <li><a href="/wiki/Evolution_of_flagella" title="Evolution of flagella">Flagella</a></li> <li><a href="/wiki/Eukaryote#Origin_of_eukaryotes" title="Eukaryote">Eukaryotes</a> <ul><li><a href="/wiki/Symbiogenesis" title="Symbiogenesis">symbiogenesis</a></li> <li><a href="/wiki/Chromosome#Eukaryotes" title="Chromosome">chromosome</a></li> <li><a href="/wiki/Endomembrane_system#Evolution" title="Endomembrane system">endomembrane system</a></li> <li><a href="/wiki/Mitochondrion#Origin_and_evolution" title="Mitochondrion">mitochondria</a></li> <li><a href="/wiki/Cell_nucleus#Evolution" title="Cell nucleus">nucleus</a></li> <li><a href="/wiki/Plastid#Origin" title="Plastid">plastids</a></li></ul></li> <li>In animals <ul><li><a href="/wiki/Evolution_of_the_eye" title="Evolution of the eye">eye</a></li> <li><a href="/wiki/Hair#Evolution" title="Hair">hair</a></li> <li><a href="/wiki/Evolution_of_mammalian_auditory_ossicles" title="Evolution of mammalian auditory ossicles">auditory ossicle</a></li> <li><a href="/wiki/Evolution_of_nervous_systems" title="Evolution of nervous systems">nervous system</a></li> <li><a href="/wiki/Evolution_of_the_brain" title="Evolution of the brain">brain</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Biological_process" title="Biological process">processes</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_ageing" title="Evolution of ageing">Aging</a> <ul><li><a href="/wiki/Death#Evolution_of_aging_and_mortality" title="Death">Death</a></li> <li><a href="/wiki/Programmed_cell_death#Evolutionary_origin_of_mitochondrial_apoptosis" title="Programmed cell death">Programmed cell death</a></li></ul></li> <li><a href="/wiki/Origin_of_avian_flight" title="Origin of avian flight">Avian flight</a></li> <li><a href="/wiki/Evolution_of_biological_complexity" title="Evolution of biological complexity">Biological complexity</a></li> <li><a href="/wiki/Cooperation_(evolution)" title="Cooperation (evolution)">Cooperation</a></li> <li><a href="/wiki/Evolution_of_color_vision" title="Evolution of color vision">Color vision</a> <ul><li><a href="/wiki/Evolution_of_color_vision_in_primates" title="Evolution of color vision in primates">in primates</a></li></ul></li> <li><a href="/wiki/Evolution_of_emotion" title="Evolution of emotion">Emotion</a></li> <li><a href="/wiki/Empathy#Evolution_across_species" title="Empathy">Empathy</a></li> <li><a href="/wiki/Evolutionary_ethics" title="Evolutionary ethics">Ethics</a></li> <li><a href="/wiki/Evolution_of_eusociality" title="Evolution of eusociality">Eusociality</a></li> <li><a href="/wiki/Immune_system#Evolution_and_other_mechanisms" title="Immune system">Immune system</a></li> <li><a href="/wiki/Metabolism#Evolution" title="Metabolism">Metabolism</a></li> <li><a href="/wiki/Monogamy_in_animals" title="Monogamy in animals">Monogamy</a></li> <li><a href="/wiki/Evolution_of_morality" title="Evolution of morality">Morality</a></li> <li><a href="/wiki/Mosaic_evolution" title="Mosaic evolution">Mosaic evolution</a></li> <li><a href="/wiki/Multicellular_organism#Evolutionary_history" title="Multicellular organism">Multicellularity</a></li> <li><a href="/wiki/Evolution_of_sexual_reproduction" title="Evolution of sexual reproduction">Sexual reproduction</a> <ul><li><a href="/wiki/Anisogamy#Evolution" title="Anisogamy">Gamete differentiation/sexes</a></li> <li><a href="/wiki/Biological_life_cycle" title="Biological life cycle">Life cycles/nuclear phases</a></li> <li><a href="/wiki/Mating_type" title="Mating type">Mating types</a></li> <li><a href="/wiki/Origin_and_function_of_meiosis" title="Origin and function of meiosis">Meiosis</a></li> <li><a href="/wiki/Sex-determination_system#Evolution" title="Sex-determination system">Sex-determination</a></li></ul></li> <li><a href="/wiki/Evolution_of_snake_venom" title="Evolution of snake venom">Snake venom</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Tempo_and_Mode_in_Evolution" title="Tempo and Mode in Evolution">Tempo and modes</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Phyletic_gradualism" title="Phyletic gradualism">Gradualism</a>/<a href="/wiki/Punctuated_equilibrium" title="Punctuated equilibrium">Punctuated equilibrium</a>/<a href="/wiki/Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li> <li><a href="/wiki/Point_mutation" title="Point mutation">Micromutation</a>/<a href="/wiki/Mutationism#Later_mutationist_theories" title="Mutationism">Macromutation</a></li> <li><a href="/wiki/Uniformitarianism" title="Uniformitarianism">Uniformitarianism</a>/<a href="/wiki/Catastrophism" title="Catastrophism">Catastrophism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Speciation" title="Speciation">Speciation</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Allopatric_speciation" title="Allopatric speciation">Allopatric</a></li> <li><a href="/wiki/Anagenesis" title="Anagenesis">Anagenesis</a></li> <li><a href="/wiki/Catagenesis_(biology)" title="Catagenesis (biology)">Catagenesis</a></li> <li><a href="/wiki/Cladogenesis" title="Cladogenesis">Cladogenesis</a></li> <li><a href="/wiki/Cospeciation" title="Cospeciation">Cospeciation</a></li> <li><a href="/wiki/Ecological_speciation" title="Ecological speciation">Ecological</a></li> <li><a href="/wiki/Hybrid_speciation" title="Hybrid speciation">Hybrid</a></li> <li><a href="/wiki/Nonecological_speciation" title="Nonecological speciation">Non-ecological</a></li> <li><a href="/wiki/Parapatric_speciation" title="Parapatric speciation">Parapatric</a></li> <li><a href="/wiki/Peripatric_speciation" title="Peripatric speciation">Peripatric</a></li> <li><a href="/wiki/Reinforcement_(speciation)" title="Reinforcement (speciation)">Reinforcement</a></li> <li><a href="/wiki/Sympatric_speciation" title="Sympatric speciation">Sympatric</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">History</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolutionary_ideas_of_the_Renaissance_and_Enlightenment" title="Evolutionary ideas of the Renaissance and Enlightenment">Renaissance and Enlightenment</a></li> <li><a href="/wiki/Transmutation_of_species" title="Transmutation of species">Transmutation of species</a></li> <li><a href="/wiki/David_Hume" title="David Hume">David Hume</a> <ul><li><i><a href="/wiki/Dialogues_Concerning_Natural_Religion" title="Dialogues Concerning Natural Religion">Dialogues Concerning Natural Religion</a></i></li></ul></li> <li><a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a> <ul><li><i><a href="/wiki/On_the_Origin_of_Species" title="On the Origin of Species">On the Origin of Species</a></i></li></ul></li> <li><a href="/wiki/History_of_paleontology" title="History of paleontology">History of paleontology</a></li> <li><a href="/wiki/Transitional_fossil" title="Transitional fossil">Transitional fossil</a></li> <li><a href="/wiki/Blending_inheritance" title="Blending inheritance">Blending inheritance</a></li> <li><a href="/wiki/Mendelian_inheritance" title="Mendelian inheritance">Mendelian inheritance</a></li> <li><a href="/wiki/The_eclipse_of_Darwinism" title="The eclipse of Darwinism">The eclipse of Darwinism</a></li> <li><a href="/wiki/Neo-Darwinism" title="Neo-Darwinism">Neo-Darwinism</a></li> <li><a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis</a></li> <li><a href="/wiki/History_of_molecular_evolution" title="History of molecular evolution">History of molecular evolution</a></li> <li><a href="/wiki/Extended_evolutionary_synthesis" title="Extended evolutionary synthesis">Extended evolutionary synthesis</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Philosophy</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Darwinism" title="Darwinism">Darwinism</a></li> <li><a href="/wiki/Alternatives_to_Darwinian_evolution" title="Alternatives to Darwinian evolution">Alternatives</a> <ul><li><a href="/wiki/Catastrophism" title="Catastrophism">Catastrophism</a></li> <li><a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a></li> <li><a href="/wiki/Orthogenesis" title="Orthogenesis">Orthogenesis</a></li> <li><a href="/wiki/Mutationism" title="Mutationism">Mutationism</a></li> <li><a href="/wiki/Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li> <li><a href="/wiki/Structuralism_(biology)" title="Structuralism (biology)">Structuralism</a> <ul><li><a href="/wiki/Spandrel_(biology)" title="Spandrel (biology)">Spandrel</a></li></ul></li> <li><a href="/wiki/Theistic_evolution" title="Theistic evolution">Theistic</a></li> <li><a href="/wiki/Vitalism" title="Vitalism">Vitalism</a></li></ul></li> <li><a href="/wiki/Teleology_in_biology" title="Teleology in biology">Teleology in biology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biogeography" title="Biogeography">Biogeography</a></li> <li><a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological genetics</a></li> <li><a href="/wiki/Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li> <li><a href="/wiki/Group_selection" title="Group selection">Group selection</a> <ul><li><a href="/wiki/Cultural_evolution" title="Cultural evolution">Cultural evolution</a></li> <li><a href="/wiki/Cultural_group_selection" title="Cultural group selection">Cultural group selection</a></li> <li><a href="/wiki/Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li></ul></li> <li><a href="/wiki/Hologenome_theory_of_evolution" title="Hologenome theory of evolution">Hologenome theory of evolution</a></li> <li><a href="/wiki/Missing_heritability_problem" title="Missing heritability problem">Missing heritability problem</a></li> <li><a href="/wiki/Molecular_evolution" title="Molecular evolution">Molecular evolution</a></li> <li><a href="/wiki/Astrobiology" title="Astrobiology">Astrobiology</a></li> <li><a href="/wiki/Phylogenetics" title="Phylogenetics">Phylogenetics</a> <ul><li><a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">Tree</a></li></ul></li> <li><a href="/wiki/Polymorphism_(biology)" title="Polymorphism (biology)">Polymorphism</a></li> <li><a href="/wiki/Protocell" title="Protocell">Protocell</a></li> <li><a href="/wiki/Systematics" title="Systematics">Systematics</a></li> <li><a href="/wiki/Transgenerational_epigenetic_inheritance" title="Transgenerational epigenetic inheritance">Transgenerational epigenetic inheritance</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <a href="/wiki/Category:Evolutionary_biology" title="Category:Evolutionary biology">Category</a></li> <li><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Symbol_portal_class.svg" class="mw-file-description" title="Portal"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/16px-Symbol_portal_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/23px-Symbol_portal_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/31px-Symbol_portal_class.svg.png 2x" data-file-width="180" data-file-height="185" /></a></span> <a href="/wiki/Portal:Evolutionary_biology" title="Portal:Evolutionary biology">Portal</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Evolutionary_psychology" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Evolutionary_psychology" title="Template:Evolutionary psychology"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Evolutionary_psychology" title="Template talk:Evolutionary psychology"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Evolutionary_psychology" title="Special:EditPage/Template:Evolutionary psychology"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Evolutionary_psychology" style="font-size:114%;margin:0 4em"><a href="/wiki/Evolutionary_psychology" title="Evolutionary psychology">Evolutionary psychology</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/wiki/History_of_evolutionary_psychology" title="History of evolutionary psychology">History</a> <ul><li><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">Evolutionary thought</a></li></ul></li> <li><a href="/wiki/Theoretical_foundations_of_evolutionary_psychology" title="Theoretical foundations of evolutionary psychology">Theoretical foundations</a> <ul><li><a href="/wiki/Adaptationism" title="Adaptationism">Adaptationism</a></li> <li><a href="/wiki/Cognitive_revolution" title="Cognitive revolution">Cognitive revolution</a></li> <li><a href="/wiki/Cognitivism_(psychology)" title="Cognitivism (psychology)">Cognitivism</a></li> <li><a href="/wiki/Gene-centered_view_of_evolution" title="Gene-centered view of evolution">Gene selection theory</a></li> <li><a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis</a></li></ul></li> <li><a href="/wiki/Criticism_of_evolutionary_psychology" title="Criticism of evolutionary psychology">Criticism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:left;"><a href="/wiki/Human_evolution" title="Human evolution">Evolutionary<br />processes</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Psychological_adaptation" title="Psychological adaptation">Adaptations</a></li> <li><a href="/wiki/Altruism_(biology)" title="Altruism (biology)">Altruism</a> <ul><li><a class="mw-selflink selflink">Cheating</a></li> <li><a href="/wiki/Hamiltonian_spite" title="Hamiltonian spite">Hamiltonian spite</a></li> <li><a href="/wiki/Reciprocal_altruism_in_humans" title="Reciprocal altruism in humans">Reciprocal</a></li></ul></li> <li><a href="/wiki/Baldwin_effect" title="Baldwin effect">Baldwin effect</a></li> <li><a href="/wiki/Spandrel_(biology)" title="Spandrel (biology)">By-products</a></li> <li><a href="/wiki/Evolutionarily_stable_strategy" title="Evolutionarily stable strategy">Evolutionarily stable strategy</a></li> <li><a href="/wiki/Exaptation" title="Exaptation">Exaptation</a></li> <li><a href="/wiki/Fitness_(biology)" title="Fitness (biology)">Fitness</a> <ul><li><a href="/wiki/Inclusive_fitness_in_humans" title="Inclusive fitness in humans">Inclusive</a></li></ul></li> <li><a href="/wiki/Kin_selection" title="Kin selection">Kin selection</a></li> <li><a href="/wiki/Evolutionary_mismatch" title="Evolutionary mismatch">Mismatch</a></li> <li><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></li> <li><a href="/wiki/Parental_investment" title="Parental investment">Parental investment</a> <ul><li><a href="/wiki/Parent%E2%80%93offspring_conflict" title="Parent–offspring conflict">Parent–offspring conflict</a></li></ul></li> <li><a href="/wiki/Sexual_selection_in_humans" title="Sexual selection in humans">Sexual selection</a> <ul><li><a href="/wiki/Costly_signaling_theory_in_evolutionary_psychology" title="Costly signaling theory in evolutionary psychology">Costly signaling</a></li> <li><a href="/wiki/Male_intrasexual_competition" title="Male intrasexual competition">Male</a>/<a href="/wiki/Female_intrasexual_competition" title="Female intrasexual competition">female intrasexual competition</a></li> <li><a href="/wiki/Mate_choice" title="Mate choice">Mate choice</a></li> <li><a href="/wiki/Sexual_dimorphism" title="Sexual dimorphism">Sexual dimorphism</a></li></ul></li> <li><a href="/wiki/Social_selection" title="Social selection">Social selection</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:left;">Areas</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolution_of_cognition" title="Evolution of cognition">Cognition</a> /<br /><a href="/wiki/Evolution_of_emotion" title="Evolution of emotion">Emotion</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Affect_(psychology)" title="Affect (psychology)">Affect</a> <ul><li><a href="/wiki/Affect_display" title="Affect display">Display</a></li> <li><a href="/wiki/Display_rules" title="Display rules">Display rules</a></li> <li><a href="/wiki/Facial_expression" title="Facial expression">Facial expression</a></li></ul></li> <li><a href="/wiki/Behavioral_modernity" title="Behavioral modernity">Behavioral modernity</a></li> <li><a href="/wiki/Cognitive_module" title="Cognitive module">Cognitive module</a>/<a href="/wiki/Modularity_of_mind" title="Modularity of mind">modularity of mind</a> <ul><li><a href="/wiki/Automatic_and_controlled_processes" title="Automatic and controlled processes">Automatic and controlled processes</a></li> <li><a href="/wiki/Computational_theory_of_mind" title="Computational theory of mind">Computational theory of mind</a></li> <li><a href="/wiki/Domain-general_learning" title="Domain-general learning">Domain generality</a></li> <li><a href="/wiki/Domain_specificity" title="Domain specificity">Domain specificity</a></li> <li><a href="/wiki/Dual_process_theory" title="Dual process theory">Dual process theory</a></li></ul></li> <li><a href="/wiki/Cognitive_tradeoff_hypothesis" title="Cognitive tradeoff hypothesis">Cognitive tradeoff hypothesis</a></li> <li><a href="/wiki/Evolution_of_the_brain" title="Evolution of the brain">Evolution of the brain</a></li> <li><a href="/wiki/Evolution_of_nervous_systems" title="Evolution of nervous systems">Evolution of nervous systems</a></li> <li><a href="/wiki/Fight-or-flight_response" title="Fight-or-flight response">Fight-or-flight response</a> <ul><li><a href="/wiki/Arachnophobia" title="Arachnophobia">Arachnophobia</a></li> <li><a href="/wiki/Fear_of_falling" title="Fear of falling">Basophobia</a></li> <li><a href="/wiki/Ophidiophobia" title="Ophidiophobia">Ophidiophobia</a></li></ul></li> <li><a href="/wiki/Folk_biology" title="Folk biology">Folk biology</a>/<a href="/wiki/Folk_taxonomy" title="Folk taxonomy">taxonomy</a></li> <li><a href="/wiki/Folk_psychology" title="Folk psychology">Folk psychology</a>/<a href="/wiki/Theory_of_mind" title="Theory of mind">theory of mind</a></li> <li><a href="/wiki/Evolution_of_human_intelligence" title="Evolution of human intelligence">Intelligence</a> <ul><li><a href="/wiki/Flynn_effect" title="Flynn effect">Flynn effect</a></li> <li><a href="/wiki/Wason_selection_task" title="Wason selection task">Wason selection task</a></li></ul></li> <li><a href="/wiki/Motor_control" title="Motor control">Motor control</a>/<a href="/wiki/Motor_skill" title="Motor skill">skill</a></li> <li><a href="/wiki/Human_multitasking" title="Human multitasking">Multitasking</a></li> <li><a href="/wiki/Neuroscience_of_sleep" title="Neuroscience of sleep">Sleep</a></li> <li><a href="/wiki/Visual_perception" title="Visual perception">Visual perception</a> <ul><li><a href="/wiki/Evolution_of_color_vision_in_primates" title="Evolution of color vision in primates">Color vision</a></li> <li><a href="/wiki/Evolution_of_the_eye" title="Evolution of the eye">Eye</a></li> <li><a href="/wiki/Na%C3%AFve_physics" title="Naïve physics">Naïve physics</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolutionary_psychology_and_culture" title="Evolutionary psychology and culture">Culture</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolutionary_aesthetics" title="Evolutionary aesthetics">Aesthetics</a> <ul><li><a href="/wiki/Darwinian_literary_studies" title="Darwinian literary studies">Literary criticism</a></li> <li><a href="/wiki/Evolutionary_musicology" title="Evolutionary musicology">Musicology</a></li></ul></li> <li><a href="/wiki/Evolutionary_anthropology" title="Evolutionary anthropology">Anthropology</a> <ul><li><a href="/wiki/Biological_anthropology" title="Biological anthropology">Biological</a></li></ul></li> <li><a href="/wiki/Biosocial_criminology" title="Biosocial criminology">Crime</a></li> <li><a href="/wiki/Evolutionary_linguistics" title="Evolutionary linguistics">Language</a> <ul><li><a href="/wiki/Origin_of_language" title="Origin of language">Origin</a></li> <li><a href="/wiki/Evolutionary_psychology_of_language" title="Evolutionary psychology of language">Psychology</a></li> <li><a href="/wiki/Origin_of_speech" title="Origin of speech">Speech</a></li></ul></li> <li><a href="/wiki/Evolution_of_morality" title="Evolution of morality">Morality</a> <ul><li><a href="/wiki/Moral_foundations_theory" title="Moral foundations theory">Moral foundations</a></li></ul></li> <li><a href="/wiki/Evolutionary_psychology_of_religion" title="Evolutionary psychology of religion">Religion</a> <ul><li><a href="/wiki/Evolutionary_origin_of_religions" class="mw-redirect" title="Evolutionary origin of religions">Origin</a></li></ul></li> <li><a href="/wiki/Cultural_universal" title="Cultural universal">Universals</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolutionary_developmental_psychology" title="Evolutionary developmental psychology">Development</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Attachment_theory" title="Attachment theory">Attachment</a></li> <li><a href="/wiki/Human_bonding" title="Human bonding">Bonding</a></li> <li><a href="/wiki/Affectional_bond" title="Affectional bond">Affectional</a>/<a href="/wiki/Maternal_bond" title="Maternal bond">maternal</a>/<a href="/wiki/Paternal_bond" title="Paternal bond">paternal bond</a></li> <li><a href="/wiki/Maternal_deprivation" title="Maternal deprivation">Caregiver deprivation</a></li> <li><a href="/wiki/Attachment_in_children" title="Attachment in children">Childhood attachment</a></li> <li><a href="/wiki/Cinderella_effect" title="Cinderella effect">Cinderella effect</a></li> <li><a href="/wiki/Cognitive_development" title="Cognitive development">Cognitive development</a></li> <li><a href="/wiki/Evolutionary_educational_psychology" title="Evolutionary educational psychology">Education</a></li> <li><a href="/wiki/Language_acquisition" title="Language acquisition">Language acquisition</a></li> <li><a href="/wiki/Personality_development" title="Personality development">Personality development</a></li> <li><a href="/wiki/Socialization" title="Socialization">Socialization</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Ergonomics" title="Ergonomics">Human factors</a> /<br /><a href="/wiki/Evolutionary_psychiatry" title="Evolutionary psychiatry">Mental health</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cognitive_ergonomics" title="Cognitive ergonomics">Cognitive ergonomics</a> <ul><li><a href="/wiki/Computer-mediated_communication" title="Computer-mediated communication">Computer-mediated communication</a></li> <li><a href="/wiki/Engineering_psychology" title="Engineering psychology">Engineering psychology</a></li> <li><a href="/wiki/Human%E2%80%93computer_interaction" title="Human–computer interaction">Human–computer interaction</a></li> <li><a href="/wiki/Media_naturalness_theory" title="Media naturalness theory">Media naturalness theory</a></li> <li><a href="/wiki/Neuroergonomics" title="Neuroergonomics">Neuroergonomics</a></li></ul></li> <li><a href="/wiki/Evolutionary_approaches_to_depression" title="Evolutionary approaches to depression">Depression</a></li> <li><a href="/wiki/Digital_media_use_and_mental_health" title="Digital media use and mental health">Digital media use and mental health</a></li> <li><a href="/wiki/Accident-proneness" title="Accident-proneness">Hypophobia</a></li> <li><a href="/wiki/Imprinted_brain_hypothesis" title="Imprinted brain hypothesis">Imprinted brain hypothesis</a></li> <li><a href="/wiki/Mind-blindness" title="Mind-blindness">Mind-blindness</a></li> <li><a href="/wiki/Psychological_effects_of_Internet_use" title="Psychological effects of Internet use">Psychological effects of Internet use</a></li> <li><a href="/wiki/Rank_theory_of_depression" title="Rank theory of depression">Rank theory of depression</a></li> <li><a href="/wiki/Evolution_of_schizophrenia" title="Evolution of schizophrenia">Schizophrenia</a></li> <li><a href="/wiki/Screen_time" title="Screen time">Screen time</a></li> <li><a href="/wiki/Smartphones_and_pedestrian_safety" title="Smartphones and pedestrian safety">Smartphones and pedestrian safety</a></li> <li><a href="/wiki/Social_aspects_of_television" title="Social aspects of television">Social aspects of television</a></li> <li><a href="/wiki/Societal_impacts_of_cars" title="Societal impacts of cars">Societal impacts of cars</a> <ul><li><a href="/wiki/Distracted_driving" title="Distracted driving">Distracted driving</a></li> <li><a href="/wiki/Lead%E2%80%93crime_hypothesis" title="Lead–crime hypothesis">Lead–crime hypothesis</a></li> <li><a href="/wiki/Mobile_phones_and_driving_safety" title="Mobile phones and driving safety">Mobile phones and driving safety</a></li> <li><a href="/wiki/Texting_while_driving" title="Texting while driving">Texting while driving</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Sexology" title="Sexology">Sex</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Human_sexual_activity" title="Human sexual activity">Activity</a></li> <li><a href="/wiki/Attachment_in_adults" title="Attachment in adults">Adult attachment</a></li> <li><a href="/wiki/Age_disparity_in_sexual_relationships" title="Age disparity in sexual relationships">Age disparity</a></li> <li><a href="/wiki/Sexual_arousal" title="Sexual arousal">Arousal</a></li> <li><a href="/wiki/Concealed_ovulation" title="Concealed ovulation">Concealed ovulation</a></li> <li><a href="/wiki/Coolidge_effect" title="Coolidge effect">Coolidge effect</a></li> <li><a href="/wiki/Sexual_desire" title="Sexual desire">Desire</a></li> <li><a href="/wiki/Sexual_fantasy" title="Sexual fantasy">Fantasy</a></li> <li><a href="/wiki/Effects_of_hormones_on_sexual_motivation" title="Effects of hormones on sexual motivation">Hormonal motivation</a></li> <li><a href="/wiki/Sexual_jealousy" title="Sexual jealousy">Jealousy</a></li> <li><a href="/wiki/Mate_guarding_in_humans" title="Mate guarding in humans">Mate guarding</a></li> <li><a href="/wiki/Mating_preferences" title="Mating preferences">Mating preferences</a></li> <li><a href="/wiki/Human_mating_strategies" title="Human mating strategies">Mating strategies</a></li> <li><a href="/wiki/Biology_and_sexual_orientation" title="Biology and sexual orientation">Orientation</a></li> <li><a href="/wiki/Ovulatory_shift_hypothesis" title="Ovulatory shift hypothesis">Ovulatory shift hypothesis</a></li> <li><a href="/wiki/Pair_bond" title="Pair bond">Pair bond</a></li> <li><a href="/wiki/Physical_attractiveness" title="Physical attractiveness">Physical</a>/<a href="/wiki/Sexual_attraction" title="Sexual attraction">Sexual attraction</a></li> <li><a href="/wiki/Human_sexuality" title="Human sexuality">Sexuality</a>/<a href="/wiki/Human_male_sexuality" title="Human male sexuality">male</a>/<a href="/wiki/Human_female_sexuality" title="Human female sexuality">female</a></li> <li><a href="/wiki/Sexy_son_hypothesis" title="Sexy son hypothesis">Sexy son hypothesis</a></li> <li><a href="/wiki/Westermarck_effect" title="Westermarck effect">Westermarck effect</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Sex_differences_in_psychology" title="Sex differences in psychology">Sex differences</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Aggression" title="Aggression">Aggression</a></li> <li><a href="/wiki/Sex_and_gender_differences_in_autism" title="Sex and gender differences in autism">Autism</a></li> <li><a href="/wiki/Sex_differences_in_cognition" title="Sex differences in cognition">Cognition</a></li> <li><a href="/wiki/Sex_differences_in_crime" title="Sex differences in crime">Crime</a></li> <li><a href="/wiki/Sexual_division_of_labour" title="Sexual division of labour">Division of labour</a></li> <li><a href="/wiki/Sex_differences_in_emotional_intelligence" title="Sex differences in emotional intelligence">Emotional intelligence</a></li> <li><a href="/wiki/Empathising%E2%80%93systemising_theory" title="Empathising–systemising theory">Empathising–systemising theory</a></li> <li><a href="/wiki/Gender_role" title="Gender role">Gender role</a></li> <li><a href="/wiki/Sex_differences_in_intelligence" title="Sex differences in intelligence">Intelligence</a></li> <li><a href="/wiki/Sex_differences_in_memory" title="Sex differences in memory">Memory</a></li> <li><a href="/wiki/Mental_disorders_and_gender" title="Mental disorders and gender">Mental health</a></li> <li><a href="/wiki/Sex_differences_in_narcissism" title="Sex differences in narcissism">Narcissism</a></li> <li><a href="/wiki/Neuroscience_of_sex_differences" title="Neuroscience of sex differences">Neuroscience</a></li> <li><a href="/wiki/Sex_differences_in_schizophrenia" title="Sex differences in schizophrenia">Schizophrenia</a></li> <li><a href="/wiki/Substance_abuse" title="Substance abuse">Substance abuse</a></li> <li><a href="/wiki/Gender_differences_in_suicide" title="Gender differences in suicide">Suicide</a></li> <li><a href="/wiki/Variability_hypothesis" title="Variability hypothesis">Variability hypothesis</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:left;">Related subjects</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Academic disciplines</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Behavioral_economics" title="Behavioral economics">Behavioral</a>/<a href="/wiki/Evolutionary_economics" title="Evolutionary economics">evolutionary economics</a></li> <li><a href="/wiki/Behavioral_epigenetics" title="Behavioral epigenetics">Behavioral epigenetics</a>/<a href="/wiki/Behavioural_genetics" title="Behavioural genetics">genetics</a></li> <li><a href="/wiki/Affective_neuroscience" title="Affective neuroscience">Affective</a>/<a href="/wiki/Behavioral_neuroscience" title="Behavioral neuroscience">behavioral</a>/<a href="/wiki/Cognitive_neuroscience" title="Cognitive neuroscience">cognitive</a>/<a href="/wiki/Evolutionary_neuroscience" title="Evolutionary neuroscience">evolutionary neuroscience</a></li> <li><a href="/wiki/Biocultural_anthropology" title="Biocultural anthropology">Biocultural anthropology</a></li> <li><a href="/wiki/Biological_psychiatry" title="Biological psychiatry">Biological psychiatry</a></li> <li><a href="/wiki/Cognitive_psychology" title="Cognitive psychology">Cognitive psychology</a></li> <li><a href="/wiki/Cognitive_science" title="Cognitive science">Cognitive science</a></li> <li><a href="/wiki/Cross-cultural_psychology" title="Cross-cultural psychology">Cross-cultural psychology</a></li> <li><a href="/wiki/Ethology" title="Ethology">Ethology</a></li> <li><a href="/wiki/Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></li> <li><a href="/wiki/Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li> <li><a href="/wiki/Functional_psychology" title="Functional psychology">Functional psychology</a></li> <li><a href="/wiki/Neuropsychology" title="Neuropsychology">Neuropsychology</a></li> <li><a href="/wiki/Philosophy_of_mind" title="Philosophy of mind">Philosophy of mind</a></li> <li><a href="/wiki/Population_genetics" title="Population genetics">Population genetics</a></li> <li><a href="/wiki/Primatology" title="Primatology">Primatology</a></li> <li><a href="/wiki/Sociobiology" title="Sociobiology">Sociobiology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Research topics</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cultural_evolution" title="Cultural evolution">Cultural evolution</a></li> <li><a href="/wiki/Evolutionary_epistemology" title="Evolutionary epistemology">Evolutionary epistemology</a></li> <li><a href="/wiki/Great_ape_language" title="Great ape language">Great ape language</a></li> <li><a href="/wiki/Human%E2%80%93animal_communication" title="Human–animal communication">Human–animal communication</a></li> <li><a href="/wiki/Missing_heritability_problem" title="Missing heritability problem">Missing heritability problem</a></li> <li><a href="/wiki/Primate_cognition" title="Primate cognition">Primate cognition</a></li> <li><a href="/wiki/Unit_of_selection" title="Unit of selection">Unit of selection</a> <ul><li><a href="/wiki/Coevolution" title="Coevolution">Coevolution</a></li> <li><a href="/wiki/Cultural_group_selection" title="Cultural group selection">Cultural group selection</a></li> <li><a href="/wiki/Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li> <li><a href="/wiki/Fisher%27s_principle" title="Fisher's principle">Fisher's principle</a></li> <li><a href="/wiki/Group_selection" title="Group selection">Group selection</a></li> <li><a href="/wiki/Hologenome_theory_of_evolution" title="Hologenome theory of evolution">Hologenome theory</a></li> <li><a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a></li> <li><a href="/wiki/Population" title="Population">Population</a></li> <li><a href="/wiki/Punctuated_equilibrium" title="Punctuated equilibrium">Punctuated equilibrium</a></li> <li><a href="/wiki/Recent_human_evolution" title="Recent human evolution">Recent human evolution</a></li> <li><a href="/wiki/Species" title="Species">Species</a></li> <li><a href="/wiki/Species_complex" title="Species complex">Species complex</a></li> <li><a href="/wiki/Transgenerational_epigenetic_inheritance" title="Transgenerational epigenetic inheritance">Transgenerational epigenetic inheritance</a></li> <li><a href="/wiki/Trivers%E2%80%93Willard_hypothesis" title="Trivers–Willard hypothesis">Trivers–Willard hypothesis</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Theoretical positions</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cultural_selection_theory" title="Cultural selection theory">Cultural selection theory</a></li> <li><a href="/wiki/Determinism" title="Determinism">Determinism</a>/<a href="/wiki/Indeterminism" title="Indeterminism">indeterminism</a> <ul><li><a href="/wiki/Biological_determinism" title="Biological determinism">Biological determinism</a></li> <li><a href="/wiki/Connectionism" title="Connectionism">Connectionism</a></li> <li><a href="/wiki/Cultural_determinism" title="Cultural determinism">Cultural determinism</a></li> <li><a href="/wiki/Environmental_determinism" title="Environmental determinism">Environmental determinism</a></li> <li><a href="/wiki/Nature_versus_nurture" title="Nature versus nurture">Nature versus nurture</a></li> <li><a href="/wiki/Psychological_nativism" title="Psychological nativism">Psychological nativism</a></li> <li><a href="/wiki/Social_constructionism" title="Social constructionism">Social constructionism</a></li> <li><a href="/wiki/Social_determinism" title="Social determinism">Social determinism</a></li> <li><a href="/wiki/Standard_social_science_model" title="Standard social science model">Standard social science model</a></li></ul></li> <li><a href="/wiki/Functionalism_(philosophy_of_mind)" title="Functionalism (philosophy of mind)">Functionalism</a></li> <li><a href="/wiki/Memetics" title="Memetics">Memetics</a></li> <li><a href="/wiki/Multilineal_evolution" title="Multilineal evolution">Multilineal evolution</a></li> <li><a href="/wiki/Neo-Darwinism" title="Neo-Darwinism">Neo-Darwinism</a></li> <li><a href="/wiki/Neoevolutionism" title="Neoevolutionism">Neoevolutionism</a></li> <li><a href="/wiki/Sociocultural_evolution" title="Sociocultural evolution">Sociocultural evolution</a></li> <li><a href="/wiki/Unilineal_evolution" title="Unilineal evolution">Unilineal evolution</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><b><a href="/wiki/List_of_evolutionary_psychologists" title="List of evolutionary psychologists">Evolutionary psychologists</a></b></li> <li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" 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