Draft:Sensory Drive

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{{Short description|Sensory drive hypothesis and examples}}{{Short description|Sensory drive hypothesis and examples}}
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The '''sensory drive hypothesis''' posits that when local environmental conditions differ between [[Conspecific|conspecific populations]], [[Animal communication|communication systems]] will adapt to these conditions<ref name=":0">{{Cite journal |last=Boughman |first=Janette Wenrick |date=December 2002 |title=How sensory drive can promote speciation |url=http://dx.doi.org/10.1016/s0169-5347(02)02595-8 |journal=Trends in Ecology & Evolution |volume=17 |issue=12 |pages=571–577 |doi=10.1016/s0169-5347(02)02595-8 |issn=0169-5347}}</ref>. Sensory drive predicts that both communication signals and [[Perception|perceptual systems]] will [[Local adaptation|adapt to these local environmental conditions]]<ref>{{Cite journal |last=West-Eberhard |first=Mary Jane |date=June 1983 |title=Sexual Selection, Social Competition, and Speciation |url=http://dx.doi.org/10.1086/413215 |journal=The Quarterly Review of Biology |volume=58 |issue=2 |pages=155–183 |doi=10.1086/413215 |issn=0033-5770}}</ref><ref name=":1">{{Cite journal |last=Endler |first=John A. |date=March 1992 |title=Signals, Signal Conditions, and the Direction of Evolution |url=http://dx.doi.org/10.1086/285308 |journal=The American Naturalist |volume=139 |pages=S125–S153 |doi=10.1086/285308 |issn=0003-0147}}</ref>. Divergence will then occur based on the intensity and direction of selection on the mating signals and on the sensory systems acquiring information regarding predators, prey, and potential mates.The '''sensory drive hypothesis''' posits that when local environmental conditions differ between [[Conspecific|conspecific populations]], [[Animal communication|communication systems]] will adapt to these conditions.<ref name=":0">{{Cite journal |last=Boughman |first=Janette Wenrick |date=December 2002 |title=How sensory drive can promote speciation |url=http://dx.doi.org/10.1016/s0169-5347(02)02595-8 |journal=Trends in Ecology & Evolution |volume=17 |issue=12 |pages=571–577 |doi=10.1016/s0169-5347(02)02595-8 |issn=0169-5347}}</ref>. Sensory drive predicts that both communication signals and [[Perception|perceptual systems]] will [[Local adaptation|adapt to these local environmental conditions]]<ref>{{Cite journal |last=West-Eberhard |first=Mary Jane |date=June 1983 |title=Sexual Selection, Social Competition, and Speciation |url=http://dx.doi.org/10.1086/413215 |journal=The Quarterly Review of Biology |volume=58 |issue=2 |pages=155–183 |doi=10.1086/413215 |issn=0033-5770}}</ref><ref name=":1">{{Cite journal |last=Endler |first=John A. |date=March 1992 |title=Signals, Signal Conditions, and the Direction of Evolution |url=http://dx.doi.org/10.1086/285308 |journal=The American Naturalist |volume=139 |pages=S125–S153 |doi=10.1086/285308 |issn=0003-0147}}</ref>. Divergence will then occur based on the intensity and direction of selection on the mating signals and on the sensory systems acquiring information regarding predators, prey, and potential mates.
The sensory drive hypothesis has two primary assumptions. The first is that greater sensory stimulation results in preferences for mates with the stimulating trait, meaning exaggerated traits are expected to have greater signal value and generate more matings because they cause a stronger response from the sensory system<ref>{{Cite journal |last1=Ryan |first1=Michael J. |last2=Keddy-Hector |first2=Anne |date=March 1992 |title=Directional Patterns of Female Mate Choice and the Role of Sensory Biases |url=http://dx.doi.org/10.1086/285303 |journal=The American Naturalist |volume=139 |pages=S4–S35 |doi=10.1086/285303 |issn=0003-0147}}</ref><ref name=":1" />. In this sense, it is possible for sensory drive to contribute to the formation of [[Fisherian runaway|runaway traits]] when [[sexual selection]] is working in the same direction as the [[Sensory bias|sensory biases]]<ref>{{Cite journal |last1=Endler |first1=John A. |last2=McLellan |first2=Tracy |date=November 1988 |title=The Processes of Evolution: Toward a Newer Synthesis |url=https://www.annualreviews.org/doi/10.1146/annurev.es.19.110188.002143 |journal=Annual Review of Ecology and Systematics |language=en |volume=19 |issue=1 |pages=395–421 |doi=10.1146/annurev.es.19.110188.002143 |issn=0066-4162}}</ref><ref name=":1" />. The second assumption is that biases are relatively fixed and show limited [[Phenotypic plasticity|developmental plasticity]]<ref>{{Cite journal |last1=TENCATE |first1=C |last2=ROWE |first2=C |date=July 2007 |title=Biases in signal evolution: learning makes a difference |url=http://dx.doi.org/10.1016/j.tree.2007.03.006 |journal=Trends in Ecology & Evolution |volume=22 |issue=7 |pages=380–387 |doi=10.1016/j.tree.2007.03.006 |pmid=17379354 |issn=0169-5347}}</ref><ref>{{Cite journal |last1=Sherman |first1=Paul W. |last2=LaReesa Wolfenbarger |first2=L. |date=June 1995 |title=Genetic correlations as tests for sensory exploitation? |url=http://dx.doi.org/10.1016/s0169-5347(00)89079-5 |journal=Trends in Ecology & Evolution |volume=10 |issue=6 |pages=246–247 |doi=10.1016/s0169-5347(00)89079-5 |pmid=21237025 |issn=0169-5347}}</ref>.The sensory drive hypothesis has two primary assumptions. The first is that greater sensory stimulation results in preferences for mates with the stimulating trait, meaning exaggerated traits are expected to have greater signal value and generate more matings because they cause a stronger response from the sensory system<ref>{{Cite journal |last1=Ryan |first1=Michael J. |last2=Keddy-Hector |first2=Anne |date=March 1992 |title=Directional Patterns of Female Mate Choice and the Role of Sensory Biases |url=http://dx.doi.org/10.1086/285303 |journal=The American Naturalist |volume=139 |pages=S4–S35 |doi=10.1086/285303 |issn=0003-0147}}</ref><ref name=":1" />. In this sense, it is possible for sensory drive to contribute to the formation of [[Fisherian runaway|runaway traits]] when [[sexual selection]] is working in the same direction as the [[Sensory bias|sensory biases]]<ref>{{Cite journal |last1=Endler |first1=John A. |last2=McLellan |first2=Tracy |date=November 1988 |title=The Processes of Evolution: Toward a Newer Synthesis |url=https://www.annualreviews.org/doi/10.1146/annurev.es.19.110188.002143 |journal=Annual Review of Ecology and Systematics |language=en |volume=19 |issue=1 |pages=395–421 |doi=10.1146/annurev.es.19.110188.002143 |issn=0066-4162}}</ref><ref name=":1" />. The second assumption is that biases are relatively fixed and show limited [[Phenotypic plasticity|developmental plasticity]]<ref>{{Cite journal |last1=TENCATE |first1=C |last2=ROWE |first2=C |date=July 2007 |title=Biases in signal evolution: learning makes a difference |url=http://dx.doi.org/10.1016/j.tree.2007.03.006 |journal=Trends in Ecology & Evolution |volume=22 |issue=7 |pages=380–387 |doi=10.1016/j.tree.2007.03.006 |pmid=17379354 |issn=0169-5347}}</ref><ref>{{Cite journal |last1=Sherman |first1=Paul W. |last2=LaReesa Wolfenbarger |first2=L. |date=June 1995 |title=Genetic correlations as tests for sensory exploitation? |url=http://dx.doi.org/10.1016/s0169-5347(00)89079-5 |journal=Trends in Ecology & Evolution |volume=10 |issue=6 |pages=246–247 |doi=10.1016/s0169-5347(00)89079-5 |pmid=21237025 |issn=0169-5347}}</ref>.
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Evidence for sensory drive in acoustic signaling and perception is lower than that in visual signaling. In a recent [[meta-analysis]], they determined that of the studies showing no support for sensory drive, 95% of them were testing the acoustic signal modality<ref name=":22" />. However, of the studies testing the sensory drive hypothesis in acoustic signaling, only two tested both components of sensory drive (signal and sensory)<ref name=":22" />. The majority of acoustic studies tested the signal component of sensory drive and there were many fewer studies testing the sensory component, suggesting this is an understudied modality<ref name=":22" />. Support for both components of the sensory drive hypothesis has been identified in birds, specifically ''[[Great tit|Parus major]]''<ref>{{Cite journal |last1=Mockford |first1=Emily J. |last2=Marshall |first2=Rupert C. |date=2009-06-03 |title=Effects of urban noise on song and response behaviour in great tits |url=http://dx.doi.org/10.1098/rspb.2009.0586 |journal=Proceedings of the Royal Society B: Biological Sciences |volume=276 |issue=1669 |pages=2979–2985 |doi=10.1098/rspb.2009.0586 |pmc=2817215 |issn=0962-8452}}</ref>, and frogs, specifically ''[[Concave-eared torrent frog|Amolops tormotus]]''<ref>{{Cite journal |last1=Feng |first1=Albert S. |last2=Narins |first2=Peter M. |last3=Xu |first3=Chun-He |last4=Lin |first4=Wen-Yu |last5=Yu |first5=Zu-Lin |last6=Qiu |first6=Qiang |last7=Xu |first7=Zhi-Min |last8=Shen |first8=Jun-Xian |date=March 2006 |title=Ultrasonic communication in frogs |url=http://dx.doi.org/10.1038/nature04416 |journal=Nature |volume=440 |issue=7082 |pages=333–336 |doi=10.1038/nature04416 |bibcode=2006Natur.440..333F |issn=0028-0836}}</ref>. In both cases, background noise is the driver underlying the sensory biases contributing to sensory drive and signal divergence.Evidence for sensory drive in acoustic signaling and perception is lower than that in visual signaling. In a recent [[meta-analysis]], they determined that of the studies showing no support for sensory drive, 95% of them were testing the acoustic signal modality<ref name=":22" />. However, of the studies testing the sensory drive hypothesis in acoustic signaling, only two tested both components of sensory drive (signal and sensory)<ref name=":22" />. The majority of acoustic studies tested the signal component of sensory drive and there were many fewer studies testing the sensory component, suggesting this is an understudied modality<ref name=":22" />. Support for both components of the sensory drive hypothesis has been identified in birds, specifically ''[[Great tit|Parus major]]''<ref>{{Cite journal |last1=Mockford |first1=Emily J. |last2=Marshall |first2=Rupert C. |date=2009-06-03 |title=Effects of urban noise on song and response behaviour in great tits |url=http://dx.doi.org/10.1098/rspb.2009.0586 |journal=Proceedings of the Royal Society B: Biological Sciences |volume=276 |issue=1669 |pages=2979–2985 |doi=10.1098/rspb.2009.0586 |pmc=2817215 |issn=0962-8452}}</ref>, and frogs, specifically ''[[Concave-eared torrent frog|Amolops tormotus]]''<ref>{{Cite journal |last1=Feng |first1=Albert S. |last2=Narins |first2=Peter M. |last3=Xu |first3=Chun-He |last4=Lin |first4=Wen-Yu |last5=Yu |first5=Zu-Lin |last6=Qiu |first6=Qiang |last7=Xu |first7=Zhi-Min |last8=Shen |first8=Jun-Xian |date=March 2006 |title=Ultrasonic communication in frogs |url=http://dx.doi.org/10.1038/nature04416 |journal=Nature |volume=440 |issue=7082 |pages=333–336 |doi=10.1038/nature04416 |bibcode=2006Natur.440..333F |issn=0028-0836}}</ref>. In both cases, background noise is the driver underlying the sensory biases contributing to sensory drive and signal divergence.
There is also support in acoustic systems for sensory drive contributing to [[speciation]] and the diversification of lineages. For example, the ''Paragalago zanzibaricus'' species complex shows support for speciation by sensory drive by looking at responses to [[conspecific]] and [[heterospecific]] signals and ear morphology<ref>{{Cite journal |last=Génin |first=Fabien |date=2021-05-03 |title=Speciation by Sensory Drive in the Paragalago zanzibaricus Species Complex |url=http://dx.doi.org/10.1007/s10764-021-00213-7 |journal=International Journal of Primatology |volume=42 |issue=3 |pages=478–498 |doi=10.1007/s10764-021-00213-7 |issn=0164-0291}}</ref>. Additionally, a study in tropical birds found that habitat characteristics predict acoustic divergence, suggesting sensory drive is playing a role in this divergence and potentially [[reproductive isolation]]<ref>{{Cite journal |last1=Tobias |first1=Joseph A. |last2=Aben |first2=Job |last3=Brumfield |first3=Robb T. |last4=Derryberry |first4=Elizabeth P. |last5=Halfwerk |first5=Wouter |last6=Slabbekoorn |first6=Hans |last7=Seddon |first7=Nathalie |date=2010-08-19 |title=Song Divergence by Sensory Drive in Amazonian Birds |url=https://doi.org/10.1111/j.1558-5646.2010.01067.x |journal=Evolution |pages=no |doi=10.1111/j.1558-5646.2010.01067.x |pmid=20561048 |issn=0014-3820}}</ref>.There is also support in acoustic systems for sensory drive contributing to [[speciation]] and the diversification of lineages. For example, the ''Paragalago zanzibaricus'' species complex shows support for speciation by sensory drive by looking at responses to [[conspecific]] and [[heterospecific]] signals and ear morphology<ref>{{Cite journal |last=Génin |first=Fabien |date=2021-05-03 |title=Speciation by Sensory Drive in the Paragalago zanzibaricus Species Complex |url=http://dx.doi.org/10.1007/s10764-021-00213-7 |journal=International Journal of Primatology |volume=42 |issue=3 |pages=478–498 |doi=10.1007/s10764-021-00213-7 |issn=0164-0291}}</ref>. Additionally, a study in tropical birds found that habitat characteristics predict acoustic divergence, suggesting sensory drive is playing a role in this divergence and potentially [[reproductive isolation]]<ref>{{Cite journal |last1=Tobias |first1=Joseph A. |last2=Aben |first2=Job |last3=Brumfield |first3=Robb T. |last4=Derryberry |first4=Elizabeth P. |last5=Halfwerk |first5=Wouter |last6=Slabbekoorn |first6=Hans |last7=Seddon |first7=Nathalie |date=2010-08-19 |title=Song Divergence by Sensory Drive in Amazonian Birds |url=https://doi.org/10.1111/j.1558-5646.2010.01067.x |journal=Evolution |pages=no |doi=10.1111/j.1558-5646.2010.01067.x |pmid=20561048 |issn=0014-3820}}</ref>
== Chemical Signals and Chemosensation ==== Chemical Signals and Chemosensation ==
[[Chemical signalling|Chemical signals]] are among the most prevalent signal modalities, but there is a lack of research focusing on sensory drive in this modality<ref name=":22" /><ref name=":02">{{Cite journal |last1=Yohe |first1=Laurel R |last2=Brand |first2=Philipp |date=2018-07-02 |title=Evolutionary ecology of chemosensation and its role in sensory drive |url=https://doi.org/10.1093/cz/zoy048 |journal=Current Zoology |volume=64 |issue=4 |pages=525–533 |doi=10.1093/cz/zoy048 |issn=1674-5507 |pmc=6084603 |pmid=30108633}}</ref>. This is likely driven by three main complications when studying chemosensation: complex chemical backgrounds, complex chemical signals, and complex genetic basis for chemosensation<ref name=":02" />. [[Chemical signalling|Chemical signals]] are among the most prevalent signal modalities, but there is a lack of research focusing on sensory drive in this modality<ref name=":22" /><ref name=":02">{{Cite journal |last1=Yohe |first1=Laurel R |last2=Brand |first2=Philipp |date=2018-07-02 |title=Evolutionary ecology of chemosensation and its role in sensory drive |url=https://doi.org/10.1093/cz/zoy048 |journal=Current Zoology |volume=64 |issue=4 |pages=525–533 |doi=10.1093/cz/zoy048 |issn=1674-5507 |pmc=6084603 |pmid=30108633}}</ref>. This is likely driven by three main complications when studying chemosensation: complex chemical backgrounds, complex chemical signals, and complex genetic basis for chemosensation<ref name=":02" />.
Recent work has discovered that chemical signals, similar to the other modalities discussed, are also affected by environmental conditions<ref name=":02" />. Under higher temperatures, chemical signals travel faster but also fade faster<ref name=":12">{{Cite journal |last1=Fuller |first1=Rebecca C |last2=Endler |first2=John A |date=2018-07-06 |title=A perspective on sensory drive |url=https://doi.org/10.1093/cz/zoy052 |journal=Current Zoology |volume=64 |issue=4 |pages=465–470 |doi=10.1093/cz/zoy052 |issn=1674-5507 |pmc=6084569 |pmid=30108627}}</ref>. Similarly, in nutrient rich or chemically complex environments, chemical signal detection can be greatly decreased<ref name=":12" />. These signals can also be subject to different interferences such as masking by other chemicals, disruption of the biological signaling pathway, or interference with the chemosensor<ref name=":12" />.Recent work has discovered that chemical signals, similar to the other modalities discussed, are also affected by environmental conditions.<ref name=":02" />. Under higher temperatures, chemical signals travel faster but also fade faster<ref name=":12">{{Cite journal |last1=Fuller |first1=Rebecca C |last2=Endler |first2=John A |date=2018-07-06 |title=A perspective on sensory drive |url=https://doi.org/10.1093/cz/zoy052 |journal=Current Zoology |volume=64 |issue=4 |pages=465–470 |doi=10.1093/cz/zoy052 |issn=1674-5507 |pmc=6084569 |pmid=30108627}}</ref>. Similarly, in nutrient rich or chemically complex environments, chemical signal detection can be greatly decreased<ref name=":12" />. These signals can also be subject to different interferences such as masking by other chemicals, disruption of the biological signaling pathway, or interference with the chemosensor<ref name=":12" />
The majority of support for sensory drive in chemosensation is considered a by-product of research that is not explicitly exploring sensory drive<ref name=":02" />. For example, in the ''[[Podarcis hispanicus]]'' species complex divergent environmental conditions (cold and humid vs. wet and dry) have led to divergence in the chemical composition of the signals<ref>{{Cite journal |last1=Harris |first1=D.James |last2=Sá-Sousa |first2=Paulo |date=April 2002 |title=Molecular Phylogenetics of Iberian Wall Lizards (Podarcis): Is Podarcis hispanica a Species Complex? |url=http://dx.doi.org/10.1006/mpev.2001.1079 |journal=Molecular Phylogenetics and Evolution |volume=23 |issue=1 |pages=75–81 |doi=10.1006/mpev.2001.1079 |pmid=12182404 |issn=1055-7903}}</ref>. This divergence is likely to contribute to producing more efficient and stable signals in their respective environments<ref name=":32">{{Cite journal |last1=Martín |first1=José |last2=López |first2=Pilar |date=2006-01-10 |title=Interpopulational differences in chemical composition and chemosensory recognition of femoral gland secretions of male lizards Podarcis hispanica: implications for sexual isolation in a species complex |url=http://dx.doi.org/10.1007/s00049-005-0326-4 |journal=Chemoecology |volume=16 |issue=1 |pages=31–38 |doi=10.1007/s00049-005-0326-4 |bibcode=2006Checo..16...31M |issn=0937-7409}}</ref>. These signals are used to mark male territory and attract female mates and both males and females show stronger responses to the conspecific signals from their respective environments<ref name=":32" /><ref>{{Cite journal |last1=Gabirot |first1=Marianne |last2=López |first2=Pilar |last3=Martín |first3=José |date=2012-01-11 |title=Differences in Chemical Sexual Signals May Promote Reproductive Isolation and Cryptic Speciation between Iberian Wall Lizard Populations |journal=International Journal of Evolutionary Biology |language=en |volume=2012 |pages=1–13 |doi=10.1155/2012/698520 |doi-access=free |pmid=22288019 |pmc=3263623 |issn=2090-8032}}</ref><ref>{{Cite journal |last1=Martín |first1=José |last2=Ortega |first2=Jesús |last3=López |first3=Pilar |date=2015-06-29 |title=Interpopulational Variations in Sexual Chemical Signals of Iberian Wall Lizards May Allow Maximizing Signal Efficiency under Different Climatic Conditions |journal=PLOS ONE |volume=10 |issue=6 |pages=e0131492 |doi=10.1371/journal.pone.0131492 |doi-access=free |pmid=26121693 |bibcode=2015PLoSO..1031492M |issn=1932-6203}}</ref>. While this is only a single example, this modality may be more important for sensory drive and the process of speciation than the lack of literature suggests<ref name=":02" />.The majority of support for sensory drive in chemosensation is considered a by-product of research that is not explicitly exploring sensory drive<ref name=":02" />. For example, in the ''[[Podarcis hispanicus]]'' species complex divergent environmental conditions (cold and humid vs. wet and dry) have led to divergence in the chemical composition of the signals<ref>{{Cite journal |last1=Harris |first1=D.James |last2=Sá-Sousa |first2=Paulo |date=April 2002 |title=Molecular Phylogenetics of Iberian Wall Lizards (Podarcis): Is Podarcis hispanica a Species Complex? |url=http://dx.doi.org/10.1006/mpev.2001.1079 |journal=Molecular Phylogenetics and Evolution |volume=23 |issue=1 |pages=75–81 |doi=10.1006/mpev.2001.1079 |pmid=12182404 |issn=1055-7903}}</ref>. This divergence is likely to contribute to producing more efficient and stable signals in their respective environments<ref name=":32">{{Cite journal |last1=Martín |first1=José |last2=López |first2=Pilar |date=2006-01-10 |title=Interpopulational differences in chemical composition and chemosensory recognition of femoral gland secretions of male lizards Podarcis hispanica: implications for sexual isolation in a species complex |url=http://dx.doi.org/10.1007/s00049-005-0326-4 |journal=Chemoecology |volume=16 |issue=1 |pages=31–38 |doi=10.1007/s00049-005-0326-4 |bibcode=2006Checo..16...31M |issn=0937-7409}}</ref>. These signals are used to mark male territory and attract female mates and both males and females show stronger responses to the conspecific signals from their respective environments<ref name=":32" /><ref>{{Cite journal |last1=Gabirot |first1=Marianne |last2=López |first2=Pilar |last3=Martín |first3=José |date=2012-01-11 |title=Differences in Chemical Sexual Signals May Promote Reproductive Isolation and Cryptic Speciation between Iberian Wall Lizard Populations |journal=International Journal of Evolutionary Biology |language=en |volume=2012 |pages=1–13 |doi=10.1155/2012/698520 |doi-access=free |pmid=22288019 |pmc=3263623 |issn=2090-8032}}</ref><ref>{{Cite journal |last1=Martín |first1=José |last2=Ortega |first2=Jesús |last3=López |first3=Pilar |date=2015-06-29 |title=Interpopulational Variations in Sexual Chemical Signals of Iberian Wall Lizards May Allow Maximizing Signal Efficiency under Different Climatic Conditions |journal=PLOS ONE |volume=10 |issue=6 |pages=e0131492 |doi=10.1371/journal.pone.0131492 |doi-access=free |pmid=26121693 |bibcode=2015PLoSO..1031492M |issn=1932-6203}}</ref>. While this is only a single example, this modality may be more important for sensory drive and the process of speciation than the lack of literature suggests<ref name=":02" />.

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