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Scleral appearance is not a correlate of domestication in mammals
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Abstract
Numerous hypotheses try to explain the unusual appearance of the human eye with its bright sclera and transparent conjunctiva and how it could have evolved from a dark-eyed phenotype, as is present in many non-human primates. Recently, it has been argued that pigmentation defects induced by self-domestication may have led to bright-eyed ocular phenotypes in humans and some other primate lineages, such as marmosets. However, it has never been systematically studied whether actual domesticated mammals consistently deviate from wild mammals in regard to their conjunctival pigmentation and if this trait might therefore be part of a domestication syndrome. Here, we test this idea by drawing phylogenetically informed comparisons from a photographic dataset spanning 13 domesticated mammal species and their closest living wild relatives ( n ≥ 15 photos per taxon). We did not recover significant differences in scleral appearance or irido-scleral contrast between domesticated and wild forms, suggesting that conjunctival depigmentation, unlike cutaneous pigmentation disorders, is not a general correlate of domestication. Regardless of their domestication status, macroscopically depigmented conjunctivae were observed in carnivorans and lagomorphs, whereas ungulates generally displayed darker eyes. For some taxa, we observed pronounced intraspecific variation, which should be addressed in more exhaustive future studies. Based on our dataset, we also present preliminary evidence for a general increase of conjunctival pigmentation with eye size in mammals. Our findings suggest that conjunctival depigmentation in humans is not a byproduct of self-domestication, even if we assume that our species has undergone such a process in its recent evolutionary history.
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There are no differences in scleral phenotypes between domesticated and their wild species.
“Recently, Caspar et al. [38] tested the hypothesis that the depigmentation of the sclera has been induced by domestication, but there are no differences in scleral phenotypes between domesticated and their wild species.”
From Are dark-eyed dogs favoured by humans? Domestication as a potential driver of iris colour difference between dogs and wolves (2023) · cites this paper as [38]
The Results report no statistically significant overall differences in scleral brightness or irido-scleral contrast between domesticated lineages and wild counterparts. But the Results and Discussion also identify pronounced differences in some pairs, including pigs and sheep, so the unqualified claim that there are no differences overstates the finding. Page numbers were not supplied. (LLM, conf=0.88)
Experimental — accuracy not yet measuredA recent study did not support the correlation between depigmentation of the sclera and domestication events.
“a recent study did not support the correlation between depigmentation of the sclera and domestication events [38].”
From Are dark-eyed dogs favoured by humans? Domestication as a potential driver of iris colour difference between dogs and wolves (2023) · cites this paper as [38]
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Cites
R: A Language and Environment for Statistical Computing
R Core Team (2000) · doi:10.32614/r.manuals · Cited by 355,817
NIH Image to ImageJ: 25 years of image analysis
Schneider, C. A., Rasband, W. S., & Eliceiri, K. W. (2012) · Nature Methods · doi:10.1038/nmeth.2089 · Cited by 64,638
phytools: an R package for phylogenetic comparative biology (and other things)
Revell, L. J. (2011) · Methods in Ecology and Evolution · doi:10.1111/j.2041-210x.2011.00169.x · Cited by 9,336
Upham, N. S., Esselstyn, J. A., & Jetz, W. (2019) · PLoS Biology · doi:10.1371/journal.pbio.3000494 · Cited by 1,235
Current perspectives and the future of domestication studies
Larson, G., Piperno, D. R., Allaby, R. G., Purugganan, M. D., Andersson, L., Arroyo‐Kalin, M., Barton, L., Vigueira, C. C., Denham, T. P., Dobney, K. M., et al. (2014) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1323964111 · Cited by 868
Wilkins, A. S., Wrangham, R. W., & Fitch, W. T. (2014) · Genetics · doi:10.1534/genetics.114.165423 · Cited by 751
Tomasello, M., Hare, B. A., Lehmann, H., & Call, J. (2006) · Journal of Human Evolution · doi:10.1016/j.jhevol.2006.10.001 · Cited by 494
Kobayashi, H., & Kohshima, S. (2001) · Journal of Human Evolution · doi:10.1006/jhev.2001.0468 · Cited by 447
Survival of the Friendliest: Homo sapiens Evolved via Selection for Prosociality
Hare, B. (2017) · Annual Review of Psychology · doi:10.1146/annurev-psych-010416-044201 · Cited by 444
The parameters of the porcine eyeball
Sanchez, I., Martin, R., Ussa, F., & Fernandez-Bueno, I. (2011) · Graefe's Archive for Clinical and Experimental Ophthalmology · doi:10.1007/s00417-011-1617-9 · Cited by 298
Ancient genomes revisit the ancestry of domestic and Przewalski’s horses
Gaunitz, C., Fages, A., Hanghøj, K., Albrechtsen, A., Khan, N., Schubert, M., Seguin-Orlando, A., Owens, I. J., Felkel, S., Bignon-Lau, O., et al. (2018) · Science · doi:10.1126/science.aao3297 · Cited by 298
Cieslak, M., Reißmann, M., Hofreiter, M., & Ludwig, A. (2011) · Biological reviews/Biological reviews of the Cambridge Philosophical Society · doi:10.1111/j.1469-185x.2011.00177.x · Cited by 283
The allometry and scaling of the size of vertebrate eyes
Howland, H. C., Merola, S. S., & Basarab, J. R. (2004) · Vision Research · doi:10.1016/j.visres.2004.03.023 · Cited by 238
Evolution of eye size and shape in primates
Ross, C. F., & Kirk, E. C. (2006) · Journal of Human Evolution · doi:10.1016/j.jhevol.2006.09.006 · Cited by 179
Neural crest derivatives in ocular development: Discerning the eye of the storm
Williams, A. L., & Bohnsack, B. L. (2015) · Birth Defects Research Part C Embryo Today Reviews · doi:10.1002/bdrc.21095 · Cited by 157
Eye movements of vertebrates and their relation to eye form and function
Land, M. F. (2014) · Journal of Comparative Physiology A · doi:10.1007/s00359-014-0964-5 · Cited by 157
Human Melanocyte Biology, Toxicology, and Pathology
Tolleson, W. H. (2005) · Journal of Environmental Science and Health Part C · doi:10.1080/10590500500234970 · Cited by 149
Evolution of facial muscle anatomy in dogs
Kaminski, J., Waller, B. M., Diogo, R., Hartstone-Rose, A., & Burrows, A. M. (2019) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1820653116 · Cited by 138
Genetic architecture of tameness in a rat model of animal domestication
Albert, F. W., Carlborg, O., Plyusnina, I., Besnier, F., Hedwig, D., Lautenschläger, S., Lorenz, D., McIntosh, J., Neumann, C., Richter, H., et al. (2009) · Genetics · doi:10.1534/genetics.109.102186 · Cited by 136
Pupil shapes and lens optics in the eyes of terrestrial vertebrates
Malmström, T., & Kröger, R. H. H. (2005) · Journal of Experimental Biology · doi:10.1242/jeb.01959 · Cited by 116
Evaluating the self‐domestication hypothesis of human evolution
Sánchez‐Villagra, M. R., & van Schaik, C. P. (2019) · Evolutionary Anthropology: Issues, News, and Reviews · doi:10.1002/evan.21777 · Cited by 115
Intraocular distribution of melanin in human, monkey, rabbit, minipig and dog eyes
Durairaj, C., Chastain, J. E., & Kompella, U. B. (2012) · Experimental Eye Research · doi:10.1016/j.exer.2012.03.004 · Cited by 84
Clinical signs of brachycephalic ocular syndrome in 93 dogs
Costa, J. C., Steinmetz, A., & Delgado, E. (2021) · Irish Veterinary Journal · doi:10.1186/s13620-021-00183-5 · Cited by 68
Scleral pigmentation leads to conspicuous, not cryptic, eye morphology in chimpanzees
Perea-García, J. O., Kret, M. E., Monteiro, A., & Hobaiter, C. (2019) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1911410116 · Cited by 62
Mayhew, J., & Gómez, J. (2015) · American Journal of Primatology · doi:10.1002/ajp.22411 · Cited by 48
Is color data from citizen science photographs reliable for biodiversity research?
Laitly, A., Callaghan, C., Delhey, K., & Cornwell, W. (2021) · Ecology and Evolution · doi:10.1002/ece3.7307 · Cited by 46
Conjunctival Primary Acquired Melanosis: Is It Time for a New Terminology?
Jakobiec, F. A. (2015) · American Journal of Ophthalmology · doi:10.1016/j.ajo.2015.11.003 · Cited by 42
Facial shape differences between rats selected for tame and aggressive behaviors
Singh, N., Albert, F. W., Plyusnina, I., Trut, L., Pӓӓbo, S., & Harvati, K. (2017) · PLoS ONE · doi:10.1371/journal.pone.0175043 · Cited by 42
Negro, J. J., Carmen Blázquez, M., & Galván, I. (2017) · Frontiers in Zoology · doi:10.1186/s12983-017-0243-8 · Cited by 42
Kano, F., Furuichi, T., Hashimoto, C., Krupenye, C., Leinwand, J. G., Hopper, L. M., Martin, C. F., Otsuka, R., & Tajima, T. (2021) · Evolution and Human Behavior · doi:10.1016/j.evolhumbehav.2021.12.004 · Cited by 42
Ocular pigmentation in humans, great apes, and gibbons is not suggestive of communicative functions
Caspar, K. R., Biggemann, M., Geissmann, T., & Begall, S. (2021) · Scientific Reports · doi:10.1038/s41598-021-92348-z · Cited by 38
Self-citationUeharu, H., Yoshida, S., Kikkawa, T., et al. (2016) · Journal of Anatomy · doi:10.1111/joa.12572 · Cited by 34
Kassab, A. (2011) · Anatomia Histologia Embryologia · doi:10.1111/j.1439-0264.2011.01097.x · Cited by 31
The Process of Animal Domestication
Sánchez-Villagra, M. R. (2022) · Princeton University Press eBooks · doi:10.1515/9780691217680 · Cited by 30
Neural crest cell genes and the domestication syndrome: A comparative analysis of selection
Rubio, A. O., & Summers, K. (2022) · PLoS ONE · doi:10.1371/journal.pone.0263830 · Cited by 29
Shared reproductive disruption, not neural crest or tameness, explains the domestication syndrome
Gleeson, B. T., & Wilson, L. A. B. (2023) · Proceedings of the Royal Society B Biological Sciences · doi:10.1098/rspb.2022.2464 · Cited by 29
The Influence of Maximum Running Speed on Eye Size: A Test of Leuckart's Law in Mammals
Heard-Booth, A. N., & Kirk, E. C. (2012) · The Anatomical Record · doi:10.1002/ar.22480 · Cited by 28
Isolation, purification and cultivation of conjunctival melanocytes
Hu, D., McCormick, S. A., Seedor, J. A., et al. (2007) · Experimental Eye Research · doi:10.1016/j.exer.2006.12.003 · Cited by 27
The evolutionary drivers of primate scleral coloration
Mearing, A. S., Burkart, J. M., Dunn, J. C., Street, S. E., & Koops, K. (2022) · Scientific Reports · doi:10.1038/s41598-022-18275-9 · Cited by 27
Domestication Phenotype Linked to Vocal Behavior in Marmoset Monkeys
Ghazanfar, A. A., Kelly, L. M., Takahashi, D. Y., Winters, S., Terrett, R., & Higham, J. P. (2020) · Current Biology · doi:10.1016/j.cub.2020.09.049 · Cited by 26
Perea‐García, J. O., Ramarajan, K., Kret, M. E., Hobaiter, C., & Monteiro, A. (2022) · Scientific Reports · doi:10.1038/s41598-022-20900-6 · Cited by 25
Normal Equine Ocular Anatomy and Eye Examination
Cooley, P. L. (1992) · Veterinary Clinics of North America Equine Practice · doi:10.1016/s0749-0739(17)30434-0 · Cited by 24
Molecular biology in conjunctival melanoma and the relationship to mucosal melanoma
Mikkelsen, L. H. (2020) · Acta Ophthalmologica · doi:10.1111/aos.14536 · Cited by 23
Perea-García, J. O., Danel, D. P., & Monteiro, A. (2021) · Symmetry · doi:10.3390/sym13071270 · Cited by 20
Oriá, A. P., Pinna, M. H., Almeida, D. S., et al. (2013) · Journal of Medical Primatology · doi:10.1111/jmp.12059 · Cited by 18
Chimpanzee (Pan troglodytes) gaze is conspicuous at ecologically-relevant distances
Whitham, W., Schapiro, S. J., Troscianko, J. T., & Yorzinski, J. L. (2022) · Scientific Reports · doi:10.1038/s41598-022-13273-3 · Cited by 17
White sclera is present in chimpanzees and other mammals
Clark, I. R., Lee, K. C., Poux, T., Langergraber, K. E., Mitani, J. C., Watts, D. P., REED, J. F., & Sandel, A. A. (2023) · Journal of Human Evolution · doi:10.1016/j.jhevol.2022.103322 · Cited by 16
Boissy, R. E., & Hornyak, T. J. (2006) · Wiley eBooks · doi:10.1002/9780470987100.ch4 · Cited by 12
Wild and Exotic Animal Ophthalmology
Giovanni, B. (2022) · Springer eBooks · doi:10.1007/978-3-030-71302-7 · Cited by 12
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- Rajathi S. Morphological and morphometrical studies of eyeball and retina in goats. Indian J Vet Anat. 2021;33(1):17–9.
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Konno, A., Aoki, H., Suzuki, E., Furuta, S., & Ueda, S. (2023) · Royal Society Open Science · doi:10.1098/rsos.230854 · Cited by 7
“[38] tested the hypothesis that the depigmentation of the sclera has been induced by domestication, but there are no differences in scleral phenotypes between domesticated and their wild species.”
Background
Metadata
- Authors
- Caspar, K. R., Hüttner, L., & Begall, S.
- Affiliations
- Czech University of Life Sciences Prague (CZ)
- University of Duisburg-Essen (DE)
- Corresponding author
- Caspar, Kai R.
- Venue
- Zoological Letters
- Published
- 2023-05-29
- Volume
- 9
- Issue
- 1
- Pages
- 12-12
- ISSN
- 2056-306X (ISSN-L)
- PMC
- PMC10228120
- OpenAlex
- W4378697795
- Cited by
- 8
- References
- 49
- Citations by year
- 2026: 2 · 2025: 2 · 2024: 3 · 2023: 1
- FWCI
- 2.03
- Type
- journal article
- Language
- en
- Update notices
- correction (2024-10-28)
- Open access
- gold
- doi.org
- Locations
- Zoological Letters (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
- Funders
- Heinrich-Heine-Universität Düsseldorf
- Topics
- Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Animal Behavior and Reproduction
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Cell Biology › melanin and skin pigmentation
- Life Sciences › Agricultural and Biological Sciences › Animal Science and Zoology › Human-Animal Interaction Studies
- Keywords
- domestication syndrome, scleral pigmentation, self-domestication, domestic mammals, eye size, marmoset, Carnivora, lagomorphs, ungulates, intraspecific variation
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