Why Aye‐Ayes See Blue

The capacity for cone‐mediated color vision varies among nocturnal primates. Some species are colorblind, having lost the functionality of their short‐wavelength‐sensitive‐1 (SWS1) opsin pigment gene. In other species, such as the aye‐aye (Daubentonia madagascariensis), the SWS1 gene remains intact. Recent studies focused on aye‐ayes indicate that this gene has been maintained by natural selection and that the pigment has a peak sensitivity (λ_max) of 406 nm, which is ～20 nm closer to the ultraviolet region of the spectrum than in most primates. The functional significance behind the retention and unusual λ_max of this opsin pigment is unknown, and it is perplexing given that all mammals are presumed to be colorblind in the dark. Here we comment on this puzzle and discuss recent findings on the color vision intensity thresholds of terrestrial vertebrates with comparable optics to aye‐ayes. We draw attention to the twilight activities of aye‐ayes and report that twilight is enriched in short‐wavelength (bluish) light. We also show that the intensity of twilight and full moonlight is probably sufficient to support cone‐mediated color vision. We speculate that the intact SWS1 opsin pigment gene of aye‐ayes is a crepuscular adaptation and we report on the blueness of potential visual targets, such as scent marks and the brilliant blue arils of Ravenala madagascariensis.     

Ancestral Loss of Short Wave-Sensitive Cone Visual Pigment in Lorisiform Prosimians,Contrasting with Its Strict Conservation in Other Prosimians

Mammals are basically dichromatic in color vision, possessing middle to long wave-sensitive (M/LWS) and the short wave-sensitive (SWS) cone opsins in the retina, whereas some nocturnal mammals lack functional SWS opsins. Prosimians, primitive primates consisting of three extant groups (Lorisiformes, Lemuriformes, and Tarsiiformes), include many nocturnal species. Among nocturnal prosimians, a species of lorisiforms, the greater galago (Otolemur crassicaudatus), is known to lack a functional SWS opsin gene, while lemuriforms and tarsiiforms appear to retain SWS opsins in the retina. It has not been established, however, whether the loss of SWS opsin is a universal phenomenon among lorisiforms and whether the functional SWS opsin genes of lemuriforms and tarsiiforms are under strict or relaxed selective constraint. To gain better insight into an association between nocturnality and loss of SWS function, we isolated and sequenced the SWS opsin genes from two species of lorisiforms, the slow loris (Nycticebus coucang; nocturnal) and the lesser galago (Galago senegalensis; nocturnal), and one species each of lemuriforms and tarsiiforms, the brown lemur (Eulemur fulvus; cathemeral) and the western tarsier (Tarsius bancanus; nocturnal), respectively. Our sequence analysis revealed that (1) the SWS opsin gene was disrupted in the common ancestor of galagids and lorisids and (2) the rate of nonsynonymous nucleotide substitution has been kept significantly lower than that of synonymous substitution in tarsier and lemur, demonstrating the presence of strict selective constraint on the SWS opsin genes in tarsiiforms and lemuriforms.     

Divergent selection for opsin gene variation in guppy (Poecilia reticulata) populations of Trinidad and Tobago

The guppy is known to exhibit remarkable interindividual variations in spectral sensitivity of middle to long wavelength-sensitive (M/LWS) cone photoreceptor cells. The guppy has four M/LWS-type opsin genes (LWS-1, LWS-2, LWS-3 and LWS-4) that are considered to be responsible for this sensory variation. However, the allelic variation of the opsin genes, particularly in terms of their absorption spectrum, has not been explored in wild populations. Thus, we examined nucleotide variations in the four M/LWS opsin genes as well as blue-sensitive SWS2-B and ultraviolet-sensitive SWS1 opsin genes for comparison and seven non-opsin nuclear loci as reference genes in 10 guppy populations from various light environments in Trinidad and Tobago. For the first time, we discovered a potential spectral variation (180 Ser/Ala) in LWS-1 that differed at an amino acid site known to affect the absorption spectra of opsins. Based on a coalescent simulation of the nucleotide variation of the reference genes, we showed that the interpopulation genetic differentiation of two opsin genes was significantly larger than the neutral expectation. Furthermore, this genetic differentiation was significantly related to differences in dissolved oxygen (DO) level, and it was not explained by the spatial distance between populations. The DO levels are correlated with eutrophication that possibly affects the color of aquatic environments. These results suggest that the population diversity of opsin genes is significantly driven by natural selection and that the guppy could adapt to various light environments through color vision changes.     

Evolution of opsin genes reveals a functional role of vision in the echolocating little brown bat (Myotis lucifugus)

Echolocating bats are able to orientate, navigate and forage without visual cues. To probe the role of vision in bats, we studied the visual opsin genes from the echolocating little brown bat (Myotis lucifugus). Short-wavelength sensitive (SWS1) opsin, middle/long-wavelength sensitive (M/LWS) opsin and rhodopsin cDNA sequences were identified from the Ensembl database and validated by the sequencing of genomic DNA. We retrieved the published orthologous genes from eleven additional representative species of mammals from GenBank and conducted an evolutionary analysis. We found that the M/LWS opsin and rhodopsin genes were both under strong purifying selection, whereas the SWS1 opsin gene has undergone positive selection at two amino acid sites and one lineage, though the main evolutionary force is still purifying selection. Two-ratio model of the SWS1 opsin gene revealed that the ω ratio for the little brown bat lineage was nearly three times lower than the background ratio, suggesting a much stronger functional constraint. Our relative rate tests show the little brown bat has a lower nonsynonymous substitution rate than those in other mammals (on average 32% lower) for the SWS1 opsin gene. However, no such significant differences were detected for the M/LWS opsin and rhodopsin genes. The results of the relative ratio tests are consistent with that of tests for selection, showing a history of purifying selection on the little brown bat opsin genes. These findings suggest a functional role of vision in the little brown bat despite being nocturnal and using echolocation. We speculate that this echolocating bat may be able to use visual cues to orientate, navigate and forage at night, to discriminate color under moonlight and starlight conditions, or to avoid predation by diurnal raptors.     

Molecular Evidence that Only Two Opsin Subfamilies,the Blue Light- (SWS2) and Green Light-Sensitive (RH2), Drive Color Vision in Atlantic Cod (Gadus morhua)

Teleosts show a great variety in visual opsin complement, due to both gene duplication and gene loss. The repertoire ranges from one subfamily of visual opsins (scotopic vision) including rod opsin only retinas seen in many deep-sea species to multiple subfamilies of visual opsins in some pelagic species. We have investigated the opsin repertoire of Atlantic cod (Gadus morhua) using information in the recently sequenced cod genome and found that despite cod not being a deep sea species it lacks visual subfamilies sensitive towards the most extreme parts of the light spectra representing UV and red light. Furthermore, we find that Atlantic cod has duplicated paralogs of both blue-sensitive SWS2 and green-sensitive RH2 subfamilies, with members belonging to each subfamily linked in tandem within the genome (two SWS2-, and three RH2A genes, respectively). The presence of multiple cone opsin genes indicates that there have been duplication events in the cod ancestor SWS2 and RH2 opsins producing paralogs that have been retained in Atlantic. Our results are supported by expressional analysis of cone opsins, which further revealed an ontogenetic change in the array of cone opsins expressed. These findings suggest life stage specific programs for opsin regulation which could be linked to habitat changes and available light as the larvae is transformed into an early juvenile. Altogether we provide the first molecular evidence for color vision driven by only two families of cone opsins due to gene loss in a teleost.     

Diurnality and cone photopigment polymorphism in strepsirrhines: examination of linkage in Lemur catta

Trichromatic color vision is routine among catarrhine primates, but occurs only as a variant form of color vision in some individuals in most platyrrhine genera. This arises from a fundamental difference in the organization of X-chromosome cone opsin genes in these two lineages: catarrhines have two opsin genes specifying middle- and long-wavelength-sensitive cone pigments, while platyrrhines have only a single gene. Some female platyrrhine monkeys achieve trichromacy because of a species polymorphism that allows the possibility of different opsin gene alleles on the two X-chromosomes. Recently, a similar opsin gene polymorphism was detected in some diurnal strepsirrhines, while at the same time appearing to be absent in any nocturnal genera. The aim of this study was to assess whether cone pigment polymorphism is inevitably linked to diurnality in strepsirrhines. Cone photopigments were measured in a species usually classified as diurnal, the ring-tailed lemur (Lemur catta), using electroretinogram flicker photometry, a noninvasive electrophysiological procedure. Each of 12 animals studied was found to have the same middle-wavelength cone pigment, with peak sensitivity at about 547 nm. In conjunction with earlier results, this implies that cone pigment polymorphism is unlikely to exist in this species and that, accordingly, such variation is not a consistently predictable feature of vision in diurnal strepsirrhines.     

Simultaneous Expression of UV and Violet SWS1 Opsins Expands the Visual Palette in a Group of Freshwater Snakes

Snakes are known to express a rod visual opsin and two cone opsins, only (SWS1, LWS), a reduced palette resulting from their supposedly fossorial origins. Dipsadid snakes in the genus Helicops are highly visual predators that successfully invaded freshwater habitats from ancestral terrestrial-only habitats. Here, we report the first case of multiple SWS1 visual pigments in a vertebrate, simultaneously expressed in different photoreceptors and conferring both UV and violet sensitivity to Helicops snakes. Molecular analysis and in vitro expression confirmed the presence of two functional SWS1 opsins, likely the result of recent gene duplication. Evolutionary analyses indicate that each sws1 variant has undergone different evolutionary paths with strong purifying selection acting on the UV-sensitive copy and d_N/d_S ∼1 on the violet-sensitive copy. Site-directed mutagenesis points to the functional role of a single amino acid substitution, Phe86Val, in the large spectral shift between UV and violet opsins. In addition, higher densities of photoreceptors and SWS1 cones in the ventral retina suggest improved acuity in the upper visual field possibly correlated with visually guided behaviors. The expanded visual opsin repertoire and specialized retinal architecture are likely to improve photon uptake in underwater and terrestrial environments, and provide the neural substrate for a gain in chromatic discrimination, potentially conferring unique color vision in the UV–violet range. Our findings highlight the innovative solutions undertaken by a highly specialized lineage to tackle the challenges imposed by the invasion of novel photic environments and the extraordinary diversity of evolutionary trajectories taken by visual opsin-based perception in vertebrates.     

Allelic Variation in Malawi Cichlid Opsins: A Tale of Two Genera

The role of sequence variation in the spectral tuning of color vision is well established in many systems. This includes the cichlids of Lake Victoria where sequence variation has been linked to environmental light gradients and speciation. The cichlids of Lake Malawi are a similar model for visual evolution, but the role of gene sequence variation in visual tuning between closely related species is unknown. This work describes such variation in multiple species of two rock-dwelling genera: Metriaclima and Labidochromis. Genomic DNA for seven cone opsin genes was sequenced and the structure of the opsin proteins was inferred. Retinal binding pocket polymorphisms were identified and compared to available data regarding spectral absorbance shifts. Sequence variation with known or potential effects on absorbance spectra were found in four genes: SWS1 (UV sensitive), SWS2B (violet sensitive), RH2Aβ (green sensitive), and LWS (red sensitive). Functional variation was distributed such that each genus had both a variable short-wavelength and long-wavelength sensitive opsin. This suggests spectral tuning is important at the margins of the cichlid visual spectrum. Further, there are two SWS1 opsin alleles that differ in sensitivity by 10 nm and are >2 MY divergent. One of these occurs in a haplotype block >1 kb. Potential haplotype blocks were found around the RH2 opsin loci. These data suggest that molecular diversification has resulted in functionally unique alleles and changes to the visual system. These data also suggest that opsin sequence variation tunes spectral sensitivities between closely related species and that the specific regions of spectral tuning are genus-specific.     

Parallel and Convergent Evolution of the Dim-Light Vision Gene RH1 in Bats (Order: Chiroptera)

Rhodopsin, encoded by the gene Rhodopsin (RH1), is extremely sensitive to light, and is responsible for dim-light vision. Bats are nocturnal mammals that inhabit poor light environments. Megabats (Old-World fruit bats) generally have well-developed eyes, while microbats (insectivorous bats) have developed echolocation and in general their eyes were degraded, however, dramatic differences in the eyes, and their reliance on vision, exist in this group. In this study, we examined the rod opsin gene (RH1), and compared its evolution to that of two cone opsin genes (SWS1 and M/LWS). While phylogenetic reconstruction with the cone opsin genes SWS1 and M/LWS generated a species tree in accord with expectations, the RH1 gene tree united Pteropodidae (Old-World fruit bats) and Yangochiroptera, with very high bootstrap values, suggesting the possibility of convergent evolution. The hypothesis of convergent evolution was further supported when nonsynonymous sites or amino acid sequences were used to construct phylogenies. Reconstructed RH1 sequences at internal nodes of the bat species phylogeny showed that: (1) Old-World fruit bats share an amino acid change (S270G) with the tomb bat; (2) Miniopterus share two amino acid changes (V104I, M183L) with Rhinolophoidea; (3) the amino acid replacement I123V occurred independently on four branches, and the replacements L99M, L266V and I286V occurred each on two branches. The multiple parallel amino acid replacements that occurred in the evolution of bat RH1 suggest the possibility of multiple convergences of their ecological specialization (i.e., various photic environments) during adaptation for the nocturnal lifestyle, and suggest that further attention is needed on the study of the ecology and behavior of bats.     

Bat Eyes Have Ultraviolet-Sensitive Cone Photoreceptors

Mammalian retinae have rod photoreceptors for night vision and cone photoreceptors for daylight and colour vision. For colour discrimination, most mammals possess two cone populations with two visual pigments (opsins) that have absorption maxima at short wavelengths (blue or ultraviolet light) and long wavelengths (green or red light). Microchiropteran bats, which use echolocation to navigate and forage in complete darkness, have long been considered to have pure rod retinae. Here we use opsin immunohistochemistry to show that two phyllostomid microbats, Glossophaga soricina and Carollia perspicillata, possess a significant population of cones and express two cone opsins, a shortwave-sensitive (S) opsin and a longwave-sensitive (L) opsin. A substantial population of cones expresses S opsin exclusively, whereas the other cones mostly coexpress L and S opsin. S opsin gene analysis suggests ultraviolet (UV, wavelengths <400 nm) sensitivity, and corneal electroretinogram recordings reveal an elevated sensitivity to UV light which is mediated by an S cone visual pigment. Therefore bats have retained the ancestral UV tuning of the S cone pigment. We conclude that bats have the prerequisite for daylight vision, dichromatic colour vision, and UV vision. For bats, the UV-sensitive cones may be advantageous for visual orientation at twilight, predator avoidance, and detection of UV-reflecting flowers for those that feed on nectar.     

Expression and Evolution of Short Wavelength Sensitive Opsins in Colugos: A Nocturnal Lineage That Informs Debate on Primate Origins

A nocturnal activity pattern is central to almost all hypotheses on the adaptive origins of primates. This enduring view has been challenged in recent years on the basis of variation in the opsin genes of nocturnal primates. A correspondence between the opsin genes and activity patterns of species in Euarchonta—the superordinal group that includes the orders Primates, Dermoptera (colugos), and Scandentia (treeshrews)—could prove instructive, yet the basic biology of the dermopteran visual system is practically unknown. Here we show that the eye of the Sunda colugo (Galeopterus variegatus) lacks a tapetum lucidum and has an avascular retina, and we report on the expression and spectral sensitivity of cone photopigments. We found that Sunda colugos have intact short wavelength sensitive (S-) and long wavelength sensitive (L-) opsin genes, and that both opsins are expressed in cone photoreceptors of the retina. The inferred peak spectral sensitivities are 451 and 562 nm, respectively. In line with adaptation to nocturnal vision, cone densities are low. Surprisingly, a majority of S-cones coexpress some L-opsin. We also show that the ratio of rates of nonsynonymous to synonymous substitutions of exon 1 of the S-opsin gene is indicative of purifying selection. Taken together, our results suggest that natural selection has favored a functional S-opsin in a nocturnal lineage for at least 45 million years. Accordingly, a nocturnal activity pattern remains the most likely ancestral character state of euprimates.     

Light exposure during embryonic and yolk-sac alevin development of Chinook salmon Oncorhynchus tshawytscha does not alter the spectral phenotype of photoreceptors

Colour vision is mediated by the expression of different visual pigments in photoreceptors of the vertebrate retina. Each visual pigment is a complex of a protein (opsin) and a vitamin A chromophore; alterations to either component affects visual pigment absorbance and, potentially, the visual capabilities of an animal. Many species of fish undergo changes in opsin expression during retinal development. In the case of salmonid fishes the single cone photoreceptors undergo a switch in opsin expression from SWS1 (ultraviolet sensitive) to SWS2 (blue-light sensitive) starting at the yolk-sac alevin stage, around the time when they first experience light. Whether light may initiate this event or produce a plastic response in the various photoreceptors is unknown. In this study, Chinook salmon Oncorhynchus tshawytscha were exposed to light from the embryonic (5 days prior to hatching) into the yolk sac alevin (25 days post hatching) stage and the spectral phenotype of photoreceptors assessed with respect to that of unexposed controls by in situ hybridization with opsin riboprobes. Light exposure did not change the spectral phenotype of photoreceptors, their overall morphology or spatial arrangement. These results concur with those from a variety of fish species and suggest that plasticity in photoreceptor spectral phenotype via changes in opsin expression may not be a widespread occurrence among teleosts.     

Mutational changes in S-cone opsin genes common to both nocturnal and cathemeral Aotus monkeys

Aotus is a platyrrhine primate that has been classically considered to be nocturnal. Earlier research revealed that this animal lacks a color vision capacity because, unlike all other platyrrhine monkeys, Aotus has a defect in the opsin gene that is required to produce short-wavelength sensitive (S) cone photopigment. Consequently, Aotus retains only a single type of cone photopigment. Other mammals have since been found to show similar losses and it has often been speculated that such change is in some fashion tied to nocturnality. Although most species of Aotus are indeed nocturnal, recent observations show that Aotus azarai, an owl monkey species native to portions of Argentina and Paraguay, displays a cathemeral activity pattern being active during daylight hours as frequently as during nighttime hours. We have sequenced portions of the S-cone opsin gene in A. azarai and Aotus nancymaae, the latter a typically nocturnal species. The S-cone opsin genes in both species contain the same fatal defects earlier detected for Aotus trivirgatus. On the basis of the phylogenetic relationships of these three species these results imply that Aotus must have lost a capacity for color vision early in its history and they also suggest that the absence of color vision is not compulsively linked to a nocturnal lifestyle.     

Evolution of opsin expression in birds driven by sexual selection and habitat

Theories of sexual and natural selection predict coevolution of visual perception with conspecific colour and/or the light environment animals occupy. One way to test these theories is to focus on the visual system, which can be achieved by studying the opsin-based visual pigments that mediate vision. Birds vary greatly in colour, but opsin gene coding sequences and associated visual pigment spectral sensitivities are known to be rather invariant across birds. Here, I studied expression of the four cone opsin genes (Lws, Rh2, Sws2 and Sws1) in 16 species of New World warblers (Parulidae). I found levels of opsin expression vary both across species and between the sexes. Across species, female, but not male Sws2 expression is associated with an index of sexual selection, plumage dichromatism. This fits predictions of classic sexual selection models, in which the sensory system changes in females, presumably impacting female preference, and co-evolves with male plumage. Expression of the opsins at the extremes of the light spectrum, Lws and Uvs, correlates with the inferred light environment occupied by the different species. Unlike opsin spectral tuning, regulation of opsin gene expression allows for fast adaptive evolution of the visual system in response to natural and sexual selection, and in particular, sex-specific selection pressures.     

Molecular cloning of cDNA encoding red opsin gene in the retinas of five Antarctic notothenioid fishes

Previous evidence suggested that notothenioid fish had lost red-sensitive (LWS) visual pigment and photoreceptors, but retained ultraviolet-sensitive (SWS1), blue-sensitive (SWS2), and green-sensitive (RH2) pigments. We used RT-PCR and Southern blot to isolate the LWS opsin gene in five notothenioid species. We determined full-coding LWS opsin sequences and genomic sequences. The expected peak absorbance of the LWS opsin, based on the five-sites rule that is primarily responsible for the spectral sensitivities in vertebrates, ranged from 541 to 553 nm. In Antarctic waters, light of this wavelength penetrates to dozens of meters. Thus, we conclude that notothenioids use tetrachromatic color vision in shallower waters, at least during the Antarctic summer.     

Signatures of functional constraint at aye-aye opsin genes: the potential of adaptive color vision in a nocturnal primate

While color vision perception is thought to be adaptively correlated with foraging efficiency for diurnal mammals, those that forage exclusively at night may not need color vision nor have the capacity for it. Indeed, although the basic condition for mammals is dichromacy, diverse nocturnal mammals have only monochromatic vision, resulting from functional loss of the short-wavelength sensitive opsin gene. However, many nocturnal primates maintain intact two opsin genes and thus have dichromatic capacity. The evolutionary significance of this surprising observation has not yet been elucidated. We used a molecular population genetics approach to test evolutionary hypotheses for the two intact opsin genes of the fully nocturnal aye-aye (Daubentonia madagascariensis), a highly unusual and endangered Madagascar primate. No evidence of gene degradation in either opsin gene was observed for any of 8 aye-aye individuals examined. Furthermore, levels of nucleotide diversity for opsin gene functional sites were lower than those for 15 neutrally evolving intergenic regions (>25 kb in total), which is consistent with a history of purifying selection on aye-aye opsin genes. The most likely explanation for these findings is that dichromacy is advantageous for aye-ayes despite their nocturnal activity pattern. We speculate that dichromatic nocturnal primates may be able to perceive color while foraging under moonlight conditions, and suggest that behavioral and ecological comparisons among dichromatic and monochromatic nocturnal primates will help to elucidate the specific activities for which color vision perception is advantageous.     

Ultraviolet photopigment sensitivity and ocular media transmittance in gulls,with an evolutionary perspective

Gulls (Laridae excluding Sternidae) appear to be the only shorebirds (Charadriiformes) that have a short wavelength sensitive type 1 (SWS1) cone pigment opsin tuned to ultraviolet (UV) instead of violet. However, the apparent UV-sensitivity has only been inferred indirectly, via the interpretation that the presence of cysteine at the key amino acid position 90 in the SWS1 opsin confers UV sensitivity. Unless the cornea and the lens efficiently transmit UV to the retina, gulls might in effect be similar to violet-sensitive birds in spectral sensitivity even if they have an ultraviolet sensitive (UVS) SWS1 visual pigment. We report that the spectral transmission of the cornea and lens of great black-backed Larus marinus and herring gulls L. argentatus allow UV-sensitivity, having a λ_T0.5 value, 344 nm, similar to the ocular media of UV sensitive birds. By molecular sequencing of the second α-helical transmembrane region of the SWS1 opsin gene we could also infer that 15 herring gulls and 16 yellow-legged gulls L. michahellis, all base-pair identical, are genetically UV-sensitive.     

Population variation in opsin expression in the bluefin killifish,<Emphasis Type="Italic"> Lucania goodei</Emphasis>: a real-time PCR study

Quantitative genetics have not been used in vision studies because of the difficulty of objectively measuring large numbers of individuals. Here, we examine the effectiveness of a molecular technique, real-time PCR, as an inference of visual components in the bluefin killifish, Lucania goodei, to determine whether there is population variation in opsin expression. Previous work has shown that spring animals possess a higher frequency of UV and violet cones and a lower frequency of yellow and red cones than swamp animals. Here, we found a good qualitative match between the population differences in opsin expression and those found previously in cone frequency. Spring animals expressed higher amounts of SWS1 and SWS2B opsins (which correspond to UV and violet photopigments) and lower amounts of RH2 and LWS opsins (which correspond to yellow and red photopigments) than swamp animals. The counterintuitive pattern between color pattern, lighting environment, and vision remains. Males with blue anal fins are more abundant in swamps where animals express fewer SWS1 and SWS2B opsins and where transmission of UV/blue wavelengths is low. Understanding this system requires quantitative genetic studies. Real-time PCR is an effective tool for studies requiring inferences of visual physiology in large numbers of individuals.Abbreviations ERG electroretinogram - MSP microspectrophotometry     

Retinal Cone Photoreceptors of the Deer Mouse Peromyscus maniculatus: Development,Topography, Opsin Expression and Spectral Tuning

A quantitative analysis of photoreceptor properties was performed in the retina of the nocturnal deer mouse, Peromyscus maniculatus, using pigmented (wildtype) and albino animals. The aim was to establish whether the deer mouse is a more suitable model species than the house mouse for photoreceptor studies, and whether oculocutaneous albinism affects its photoreceptor properties. In retinal flatmounts, cone photoreceptors were identified by opsin immunostaining, and their numbers, spectral types, and distributions across the retina were determined. Rod photoreceptors were counted using differential interference contrast microscopy. Pigmented P. maniculatus have a rod-dominated retina with rod densities of about 450.000/mm² and cone densities of 3000 - 6500/mm². Two cone opsins, shortwave sensitive (S) and middle-to-longwave sensitive (M), are present and expressed in distinct cone types. Partial sequencing of the S opsin gene strongly supports UV sensitivity of the S cone visual pigment. The S cones constitute a 5-15% minority of the cones. Different from house mouse, S and M cone distributions do not have dorsoventral gradients, and coexpression of both opsins in single cones is exceptional (<2% of the cones). In albino P. maniculatus, rod densities are reduced by approximately 40% (270.000/mm²). Overall, cone density and the density of cones exclusively expressing S opsin are not significantly different from pigmented P. maniculatus. However, in albino retinas S opsin is coexpressed with M opsin in 60-90% of the cones and therefore the population of cones expressing only M opsin is significantly reduced to 5-25%. In conclusion, deer mouse cone properties largely conform to the general mammalian pattern, hence the deer mouse may be better suited than the house mouse for the study of certain basic cone properties, including the effects of albinism on cone opsin expression.