· How many distinct molecular paths lead to the same phenotype? One approach to this question has been to examine the genetic basis of convergent traits. “I’m using a compact point and shoot digital camera and i would like to ask it is worth it to upgrade to a DSLR camera? How huge a difference do DSLR cameras make.The Convergent Evolution of Blue Iris Pigmentation in Primates Took Distinct Molecular Paths. In natural populations, the number of possible genetic changes leading to a given trait is determined by the combination of two factors: the number of loci at which mutations influence the phenotype (the mutational target size) and the proportion of these mutations that do not have prohibitively deleterious consequences on other phenotypes (the extent of pleiotropy; Stern and Orgogozo, 2. Stern, 2. 01. 1). Examples of convergent evolution, the independent acquisition of the same trait in different populations or species, may be highly informative with regard to questions of evolutionary predictability and constraint because these likely represent multiple solutions to similar selective pressures (see, e. Conway Morris, 2. Christin et al., 2. Losos, 2. 01. 1). To date, cases of convergence have been found to occur through independent coding changes in the same gene (e. Mundy, 2. 00. 5; Christin et al., 2. Gross et al., 2. 00. Kingsley et al., 2. Miller et al., 2. Tishkoff et al., 2. Aminetzach et al., 2. Feldman et al., 2. Li et al., 2. 01. Liu et al., 2. 01. Shen et al., 2. 01. In other cases, changes in distinct loci result in the evolution of the same trait (e. ![]() Chen et al., 1. 99. Hoekstra et al., 2. Borowsky, 2. 00. 8; Steiner et al., 2. Most studies have found that convergent phenotypes tend to evolve through changes within the same gene (Conte et al., 2. Martin and Orgogozo, in press) or within a particular pathway (e. Chan et al., 2. 01. Tenaillon et al., 2. Many of these reported cases of phenotypic convergence involve pigmentation (see Hubbard et al., 2. Miller et al., 2. Gross et al., 2. 00. Hoekstra et al., 2. Kingsley et al., 2. Steiner et al., 2. Mundy, 2. 00. 5). In primates, blue iris pigmentation has been documented in four different lineages. In humans (Homo sapiens), the prevalence of the trait increases with latitude from 2. European populations (Zanetti et al., 1. Laeng et al., 2. 00. European admixture (Frudakis et al., 2. In Japanese macaques (Macaca fuscata), which are endemic to the broad- leaved deciduous and evergreen forests of Japan (Oi, 2. Abe et al., 2. 00. Shodoshima and Kyushu islands (Yamagiwa, 1. Zhang and Watanabe, 2. Blue irises have also been reported in the brown spider monkey (Ateles hybridus, formerly Ateles belzebuth hybridus; Hernandez- Camacho and Cooper, 1. Konstant et al., 1. Colombian black spider monkey (Ateles geoffroyi or fusciceps, subspecies rufiventris; Hernandez- Camacho and Cooper, 1. Defler et al., 2. Colombia, Panama, and Venezuela (Mondolfi and Eisenberg, 1. Defler et al., 2. Although the prevalence of this trait in these species has not been extensively documented, approximately 1. Colombia has blue irises (R. Rimbach and A. Link, personal communication). In contrast to these other species, all blue- eyed black lemurs (Eulemur flavifrons, formerly Eulemur macaco flavifrons) have blue irises, whereas in the most closely related species, the black lemur (Eulemur macaco, formerly Eulemur macaco macaco), all individuals have brown irises (Mittermeier et al., 2. These sister species inhabit primary and secondary tropical sub- humid forests in a narrow range in northwestern Madagascar (Rabarivola et al., 1. Andrianjakarivelo, 2. Randriatahina and Rabarivola, 2. Rabarivola et al., 1. Given that almost all other primates have brown or yellow irises (Kobayashi and Kohshima, 2. Beyond these broad strokes, the extent of iris pigmentation variation within and between species has not been characterized. In particular, it is not yet clear whether the reported blue irises represent the same phenotype in all species. Demonstrating such similarity is important, as if the derived phenotypes were different, we would anticipate the involvement of distinct genetic loci. Thus, mapping the traits would not provide additional information with regard to evolutionary constraint. Quantitative measures of iris pigmentation variation, recently demonstrated to be associated with genetic variation in humans (Liu et al., 2. Edwards et al., 2. In humans, the phenotype of blue iris pigmentation has well understood molecular and genetic underpinnings. Observable human “eye color” depends upon the amount and type of melanin within the outermost layers of the iris (Prota et al., 1. Wielgus and Sarna, 2. Whether human irises are blue or brown can be predicted with > 8. SNP), rs. 12. 91. OCA2 (Eiberg et al., 2. Sturm et al., 2. 00. Liu et al., 2. 00. Visser et al., 2. The OCA2 gene itself encodes a transport protein expressed in the iris that influences the development of melanocytes (Rinchik et al., 1. Brilliant, 2. 00. In addition to the main effect SNP, variants within OCA2 and at least 1. Kanetsky et al., 2. Frudakis et al., 2. Graf et al., 2. 00. Duffy et al., 2. 00. Kayser et al., 2. Sturm et al., 2. 00. Liu et al., 2. 00. Age strongly influences human iris color variation, and sex has a significant but weak effect (Liu et al., 2. Our understanding of the genetic basis of blue iris pigmentation in humans can inform hypotheses regarding the genetic basis in other primates. In humans, several coding sequence mutations and deletions of coding regions in OCA2 cause oculocutaneous albinism, or severely reduced pigmentation of the hair, skin, and irises (Manga and Orlow, 1. Oetting and King, 1. SNP rs. 12. 91. 38. Sulem et al., 2. 00. Branicki et al., 2. If dramatic pigmentation reduction in skin or fur were deleterious in other primates, we may expect blue irises to evolve via regulatory mutations in these species as well. Notably, SNP rs. 12. OCA2 expression. Bradley et al. They identified one variant that was polymorphic within the blue- eyed black lemurs but not the black lemurs sequenced, a pattern that could be suggestive of this variant or a linked site contributing to pigmentation differences. Several lines of evidence support the idea that blue irises have fitness effects in humans. The rarity of observation of the phenotype in nature, in combination with observed deleterious pleiotropic consequences in humans (Eagle Jr, 1. Imesch et al., 1. Metallinos et al., 1. Santschi et al., 1. Yang et al., 1. 99. Smith et al., 2. 00. Pingault et al., 2. Juraschko et al., 2. Geigy et al., 2. 00. Hauswirth et al., 2. Population genetic evidence also supports recent positive selection at the locus responsible for the majority of the phenotypic variation in humans (Voight et al., 2. Donnelly et al., 2. Yang et al., 2. 01. Although the mechanism driving this population genetic signature is unclear, direct selection for blue irises to improve short wavelength perception (Bornstein, 1. Laeng et al., 2. 00. D absorption (see, e. Jablonski and Chaplin, 2. Parra, 2. 00. 7), and sexual selection (Darwin, 1. Diamond, 1. 99. 2; Aoki, 2. Frost, 2. 00. 6; Laeng et al., 2. Evidence for an adaptive value of blue irises in non- human primates is more limited; however, pleiotropy and sexual selection may also have played a role in these species. In Japanese macaques, the presence of blue irises is moderately correlated with reduced pigment in fur (Yamagiwa, 1. Of note, Japanese macaques have the northern- most range of any non- human primate (3. Oi, 2. 00. 2), so selection either for increased absorption of UVB or for short wavelength perception could have occurred in this species. In contrast, neither blue- eyed black lemurs (which are more lightly pigmented than black lemurs among females; Mittermeier et al., 2. Several adaptive hypotheses involving blue irises, including sexual selection and species recognition, have been proposed for the blue- eyed black lemur (Bradley et al., 2. Alternatively, blue irises may have risen to high frequency by chance (i. Identifying the loci that underlie blue iris pigmentation in these species would help in evaluating hypotheses about the selective pressures shaping the evolution of this phenotype. MATERIALS AND METHODSObtaining and selecting photographs. Photographs of five black lemurs and eleven blue- eyed black lemurs from the Duke Lemur Center (DLC), Durham, North Carolina, were kindly provided by David Haring (DLC). Photographs of 4. Japanese macaques housed at the Primate Research Institute (PRI) in Inuyama, Aichi Prefecture, Japan, were taken by PRI staff. We obtained photographs of 1. Japanese macaques in the Choshikei Monkey Park on Shodoshima Island, Kagawa prefecture, Japan. Photographs of eight brown spider monkeys were kindly provided by Rebecca Rimbach at the German Primate Center, Göttingen. Photographs of 1.
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