Middle-wavelength-sensitive Research Articles

Crustacean conundrums: a review of opsin diversity and evolution.

Knowledge of crustacean vision is lacking compared to the more well-studied vertebrates and insects. While crustacean visual systems are typically conserved morphologically, the molecular components (i.e. opsins) remain understudied. This review aims to characterize opsin diversity across crustacean lineages for an integrated view of visual system evolution. Using publicly available data from 95 species, we identified opsin sequences and classified them by clade. Our analysis produced 485 putative visual opsins and 141 non-visual opsins. The visual opsins were separated into six clades: long wavelength sensitive (LWS), middle wavelength sensitive (MWS) 1 and 2, short wavelength or ultraviolet sensitive (SWS/UVS) and a clade of thecostracan opsins, with multiple LWS and MWS opsin copies observed. The SWS/UVS opsins were relatively conserved in most species. The crustacean classes Cephalocarida, Remipedia and Hexanauplia exhibited reduced visual opsin diversity compared to others, with the malacostracan decapods having the highest opsin diversity. Non-visual opsins were identified from all investigated classes except Cephalocarida. Additionally, a novel clade of non-visual crustacean-specific, R-type opsins (Rc) was discovered. This review aims to provide a framework for future research on crustacean vision, with an emphasis on the need for more work in spectral characterization and molecular analysis. This article is part of the theme issue 'Understanding colour vision: molecular, physiological, neuronal and behavioural studies in arthropods'.

Characterization and Comparison of Eye Development and Phototransduction Genes in Deep- and Shallow-Water Shrimp Alvinocaris longirostris and Palaemon carinicauda

The investigations of the molecular components of eye development and phototransduction in deep-sea species are important to elucidate the mechanism of their adaptation to dim light. In this study, eye transcriptomes of the shrimp Alvinocaris longirostris from the deep-sea chemosynthetic ecosystem and the shallow-water shrimp Palaemon carinicauda were compared. Two Pax6 homologs with low expression levels were identified in both species, which are essential transcription factors in eye development. This finding implies that the development of the two shrimp eyes at early embryo–larvae stages might be similar. The multiple components of the phototransduction pathway were identified in both species. However, the number of phototransduction components was significantly reduced in A. longirostris, as well as expression level. Particularly, short-wavelength/UV-sensitive (SWS/UVS) opsins were absent in A. longirostris and only one putative middle-wavelength-sensitive (MWS) opsin was identified in this species. The conserved sites and structures of the putative LWS opsins were found between deep-sea and shallow-water decapods, indicating that the opsins in deep-sea crustaceans may also conserve their spectral absorption and signal transduction function. Phylogenetic analyses supported the monophyly of LWS opsins and SWS/UVS opsins in arthropods, while the MWS clade fell outside of the main arthropod LWS clade. The results are expected to provide baseline for study of visual adaptation in deep-sea shrimps.

Photoreceptors and eyes of pikeperch Sander lucioperca, pike Esox lucius, perch Perca fluviatilis and roach Rutilus rutilus from a clear and a brown lake

The photoreceptors and eyes of four fish species commonly cohabiting Fennoscandian lakes with different light transmission properties were compared: pikeperch Sander lucioperca, pike Esox lucius, perch Perca fluviatilis and roach Rutilus rutilus. Each species was represented by individuals from a clear (greenish) and a humic (dark brown) lake in southern Finland: Lake Vesijärvi (LV; peak transmission around 570 nm) and Lake Tuusulanjärvi (LT; peak transmission around 630 nm). In the autumn, all species had almost purely A2-based visual pigments. Rod absorption spectra peaked at c.526 nm (S. lucioperca), c. 533 nm (E. lucius) and c. 540 nm (P. fluviatilis and R. rutilus), with no differences between the lakes. Esox lucius rods had remarkably long outer segments, 1.5-2.8-fold longer than those of the other species. All species possessed middle-wavelength-sensitive (MWS) and long-wavelength-sensitive (LWS) cone pigments in single, twin or double cones. Rutilus rutilus also had two types of short-wavelength sensitive (SWS) cones: UV-sensitive [SWS1] and blue-sensitive (SWS2) cones, although in the samples from LT no UV cones were found. No other within-species differences in photoreceptor cell complements, absorption spectra or morphologies were found between the lakes. However, E. lucius eyes had a significantly lower focal ratio in LT compared with LV, enhancing sensitivity at the expense of acuity in the dark-brown lake. Comparing species, S. lucioperca was estimated to have the highest visual sensitivity, at least two times higher than similar-sized E. lucius, thanks to the large relative size of the eye (pupil) and the presence of a reflecting tapetum behind the retina. High absolute sensitivity will give a competitive edge also in terms of short reaction times and long visual range.

Molecular Characterization of Copepod Photoreception.

Copepod crustaceans are an abundant and ecologically significant group whose basic biology is guided by numerous visually guided behaviors. These behaviors are driven by copepod eyes, including naupliar eyes and Gicklhorn's organs, which vary widely in structure and function among species. Yet little is known about the molecular aspects of copepod vision. In this study we present a general overview of the molecular aspects of copepod vision by identifying phototransduction genes from newly generated and publicly available RNA-sequencing data and assemblies from 12 taxonomically diverse copepod species. We identify a set of 10 expressed transcripts that serve as a set of target genes for future studies of copepod phototransduction. Our more detailed evolutionary analyses of the opsin gene responsible for forming visual pigments found that all of the copepod species investigated express two main groups of opsins: middle-wavelength-sensitive (MWS) opsins and pteropsins. Additionally, there is evidence from a few species (e.g., Calanus finmarchicus, Eurytemora affinis, Paracyclopina nana, and Lernaea cyprinacea) for the expression of two additional groups of opsins-the peropsins and rhodopsin 7 (Rh7) opsins-at low levels or distinct developmental stages. An ontogenetic analysis of opsin expression in Calanus finmarchicus found the expression of a single dominant MWS opsin, as well as evidence for differences in expression across development in some MWS, pteropsin, and Rh7 opsins, with expression peaking in early naupliar through early copepodite stages.

The Genetic and Evolutionary Drives behind Primate Color Vision

Primate color vision is based on two to three cone types in the retina, each expressing a different class of visual pigment, making them the only mammals that possess trichromacy. These pigment classes are the short wavelength-sensitive (SWS1) pigment and the long wavelength-sensitive (LWS) pigment, orthologues of the same pigments found in many other vertebrates, as well as the middle wavelength-sensitive (MWS) pigment, a paralogue to the LWS pigment. Trichromacy was achieved differently in Old World and New World primates. In Old World primates, a duplication of the LWS opsin gene occurred giving rise to a “red-sensitive” or L pigment and a “green-sensitive” or M pigment. Their corresponding L and M genes are adjacent on the X chromosome which, together with their high sequence homology, is the underlying cause for the high frequency of red-green color blindness seen in humans. In New World primates and prosimians, however, the mechanism leading to trichromacy, with one exception, is based on a single polymorphic LWS gene, from which different allelic variants encode pigments with differing spectral peaks. X chromosome inactivation limits expression to just one gene per photoreceptor meaning that trichromacy is only seen in females; while all male are red-green color blind. Despite several leading hypothesis, the reasons for the different evolutionary paths taken by Old and New World primates for trichromacy are still unclear and remain to be confirmed.

Ontogenetic changes in spectral sensitivity and retinal topography in the retina of the yellowtail kingfish (Seriola lalandi): Implications for the global Seriola aquaculture industry

The visual environment of several aquaculture species has been shown to be crucial for increased survival and growth. However, very little is known about the visual system of the increasingly popular Seriola genus and its developmental changes. We investigated the ontogenetic changes in colour vision and spatial resolution in the yellowtail kingfish, S. lalandi and discuss them in relation to past behavioural feeding experiments. Microspectrophotometry was used to measure the spectral sensitivities of photoreceptor visual pigments in larvae and sub-adults. The yellowtail kingfish larva has a broad spectral range including ultraviolet-sensitive (UVS) single cones, middle-wavelength-sensitive (MWS) single cones, MWS twin cones and long-wavelength-sensitive (LWS) twin cones. The overall spectral range narrows during development through the loss of the LWS twin cones and a general short-wavelength shift in the spectral sensitivities of the other photoreceptor types. As a sub-adult the yellowtail kingfish also gains a short-wave-sensitive (SWS) single cone. This suggests that different lighting conditions will be optimal for larvae and sub-adults. No changes are observed in retinal ganglion cell (RGC) distribution between juveniles and sub-adults. A temporal concentric increase in RGC density and a weak horizontal band of medium RGC density (30% of the peak density) are present at both stages. This suggests that no changes in the feeding behaviour will occur throughout growth although the sub-adult possesses 1.6 times higher spatial resolution which would increase feeding accuracy at higher pellet densities. Research into the spectral properties of feed types combined with the data obtained here will allow for better contrast between prey and background by optimising lighting conditions.

Differences in spectral sensitivity within and among species of darters (genus Etheostoma)

We examined variation in the visual system both within and among seven species of darters, colorful freshwater fishes of the genus Etheostoma. Using microspectrophotometry, we found that darters possess rod photoreceptor cells, single cone photoreceptor cells containing middle wavelength sensitive (MWS) visual pigments, and twin photoreceptor cells containing (LWS) visual pigments. No variation in peak sensitivity was detected among species or individuals in the rod class. In the MWS class, significant variation was detected among species and a strong statistical trend suggests differences among individuals. By contrast, all differences in the LWS class could be attributed to variation among individuals. Patterns of variation detected among species, among individuals, and among cone classes suggest that complex patterns of selection may be shaping the visual system of these fishes. Further, differences among individuals may have important consequences for visually based behaviors.

Estimated absorbance spectra of the visual pigments of the North Atlantic right whale (Eubalaena glacialis)

To assess the spectral sensitivities of the retinal visual pigments from the North Atlantic right whale (Eubalaena glacialis), we have cloned and sequenced two exons from the rod opsin gene and two exons from the middle-wavelength sensitive (MWS) cone opsin gene in order to determine the amino acids at positions known to be key regulators of the spectral location of the absorbance maximum (λmax). Based on previous mutagenesis models we estimate that the right whale possesses a rod visual pigment with a λmax of 499 nm and a MWS cone visual pigment with a λmax of 524 nm. Although the MWS cone visual pigment from the right whale is blue-shifted in its spectral sensitivity like those from odontocetes, the spectral sensitivity of the right whale rod visual pigment is similar to those from terrestrial mammals.

The molecular genetics and evolution of colour and polarization vision in stomatopod crustaceans

Stomatopod crustaceans have the most complex assemblage of visual receptor classes known; retinas of many species are thought to express up to 16 different visual pigments. Physiological studies indicate that stomatopods contain up to six distinct middle-wavelength-sensitive (MWS) photoreceptor classes, suggesting that no more than six different MWS opsin gene copies exist per species. However, we previously reported the unexpected expression of 6-15 different MWS genes in retinas of each of five stomatopod species (Visual Neurosci 26: 255-266, 2009). Here, we present a review of the results reported in this publication, plus new results that shed light on the origins of the diverse colour and polarization visual capabilities of stomatopod crustaceans. Using in situ hybridization of opsins in photoreceptor cells, we obtained new results that support the hypothesis of an ancient functional division separating spatial and polarizational vision from colour vision in the stomatopods. Since evolutionary trace analysis indicates that stomatopod MWS opsins have diverged both with respect to spectral tuning and to cytoplasmic interactions, we have now further analyzed these data in an attempt to uncover the origins, diversity and potential specializations among clades for specific visual functions. The presence of many clusters of highly similar transcripts suggests exuberant opsin gene duplication has occurred in the stomatopods, together with more conservative, ancient gene duplication events within the stem crustacean lineage. Phylogenetic analysis of opsin relatedness suggests that opsins specialized for colour vision have diverged from those devoted to polarization vision, and possibly motion and spatial vision.

The Green-absorbing Drosophila Rh6 Visual Pigment Contains a Blue-shifting Amino Acid Substitution That Is Conserved in Vertebrates

The molecular mechanisms that regulate invertebrate visual pigment absorption are poorly understood. Through sequence analysis and functional investigation of vertebrate visual pigments, numerous amino acid substitutions important for this adaptive process have been identified. Here we describe a serine/alanine (S/A) substitution in long wavelength-absorbing Drosophila visual pigments that occurs at a site corresponding to Ala-292 in bovine rhodopsin. This S/A substitution accounts for a 10-17-nm absorption shift in visual pigments of this class. Additionally, we demonstrate that substitution of a cysteine at the same site, as occurs in the blue-absorbing Rh5 pigment, accounts for a 4-nm shift. Substitutions at this site are the first spectrally significant amino acid changes to be identified for invertebrate pigments sensitive to visible light and are the first evidence of a conserved tuning mechanism in vertebrate and invertebrate pigments of this class.

The cone visual pigments of an Australian marsupial, the tammar wallaby (Macropus eugenii): sequence, spectral tuning, and evolution.

Studies on marsupial color vision have been limited to very few species. There is evidence from behavioral, electroretinographic (ERG), and microspectrophotometric (MSP) measurements for the existence of both dichromatic and trichromatic color vision. No studies have yet investigated the molecular mechanisms of spectral tuning in the visual pigments of marsupials. Our study is the first to determine the mRNA sequence, infer the amino acid sequence, and determine, by in vitro expression, the spectra of the cone opsins of a marsupial, the tammar wallaby (Macropus eugenii). This yielded some information on mechanisms and evolution of spectral tuning of these pigments. The tammar wallaby retina contains only short-wavelength sensitive (SWS) and middle-wavelength sensitive (MWS) pigment mRNAs. This predicts dichromatic color vision, which is consistent with conclusions from previous behavioral studies ( Hemmi 1999). We found that the wallaby has a SWS1 class pigment of 346 amino acids. Sequence comparison with eutherian SWS pigments predicts that this SWS1 pigment absorbs maximally (lambdamax) at 424 nm and, therefore, is a blue rather than a UV pigment. This (lambdamax) is close to that of the in vitro-expressed wallaby SWS pigment (lambdamax of 420 +/- 2 nm) and to that determined behaviorally (420 nm). The difference from the mouse UV pigment (lambdamax of 359 nm) is largely accounted for by the F86Y substitution, in agreement with in vitro results comparing a variety of other SWS pigments. This suggests that spectral tuning employing F86Y substitution most likely arose independently in the marsupials and ungulates as a result of convergent evolution. An apparently different mechanism of spectral tuning of the SWS1 pigments, involving five amino acid positions, evolved in primates. The wallaby MWS pigment has 363 amino acids. Species comparisons at positions critical to spectral tuning predict a lambdamax near 530 nm, which is close to that of the in vitro-expressed pigment (529 +/- 1 nm), but quite different from the value of 539 nm determined by microspectrophotometry. Introns interrupt the coding sequences of the wallaby, mouse, and human MWS pigment sequences at the same corresponding nucleotide positions. However, the length of introns varies widely among these species.

Rearing in different photic and chromatic environments modifies spectral responses of cone horizontal cells in adult fish retina.

We investigated chromatic processing in the outer retina of the cichlid fish Aequidens pulcher. Intracellular recordings from cone-specific horizontal cells (CHCs) revealed that the two morphologically identified types (H1 and H2) also have different spectral responses. H1-L cells hyperpolarize to all wavelengths ("luminosity"). H2-Cb cells depolarize to long wavelengths and hyperpolarize to short wavelengths ("chromaticity", biphasic). Furthermore, we verified by immunocytochemistry that H2-Cb cells of A. pulcher predominantly contact the middle-wavelength-sensitive (MWS) members of double cones. Developmental plasticity in the cone-CHC networks was induced by rearing fish under conditions of spectral deprivation and different levels of white light. H1-L spectral responses were unaffected by the rearing conditions. Different intensity levels of white light and deprivation of long wavelengths during rearing both induced changes in the spectral responses of H2-Cb. Deprivation of short and middle wavelengths had no effect. Our results indicate that spectral processing in the outer retina of fishes can be modulated in response to different visual experiences and suggest that developmental fine tuning of the color-vision system occurs at early levels of visual processing.

Color vision in two observers with highly biased LWS/MWS cone ratios

Two sisters, heterozygous carriers for congenital X-linked protanopia, were diagnosed as normal trichromats by the Rayleigh match on the anomaloscope. The heterozygous state was established by molecular analysis of their visual pigment genes. The normal color match establishes that the spectral sensitivities of their long-wavelength-sensitive (LWS) and middle-wavelength-sensitive (MWS) cone visual photopigments are within normal variability. Their FM 100-hue test error scores were low, demonstrating superior chromatic discrimination. Heterochromatic flicker photometric (HFP) spectral sensitivities were like those of protanopes. The estimated LWS/MWS cone ratios from the HFP data were 0.09 1 and 0.03 1 , compared with ratios in the range of 0.6 1 to 10 1 for typical normal trichromats. Measurements of chromatic grating acuity on chromatically selective backgrounds were performed to study the cone mosaic. The data were consistent with a sparsity of LWS cones. Both protan carriers showed normal spectral sensitivities for all three cone types under cone isolating chromatic adaptation and normal three-peaked curves for increment thresholds on at white pedestal. Hue estimation, run on one carrier was normal. The equilibrium yellow locus was measured in the other carrier and was in the range of normal trichromats. The data indicate that normal color vision can occur even when the LWS/MWS cone ratio is quite abnormal.

Positional information and opsin identity in retinal cones.

To test the hypothesis that local environmental cues regulate the expression of middle wavelength-sensitive (MWS) and short wavelength-sensitive (SWS) opsins in cone photoreceptors, we examined the development of the neonatal mouse retina in an organotypic culture system. The segregation of MWS and SWS cones into dorsal and ventral fields in the mouse retina offers an opportunity to isolate a phenotypically homogeneous population of immature cones prior to opsin expression. Retinae were harvested from mice ranging in age from birth (P0) to P18 and maintained in vitro for up to 4 weeks. Cones from newborn mice were first immunoreactive to SWS opsin-specific antibodies (OS-2 and JH455) after 5 days in vitro, which corresponds to a time course similar to that in vivo. The topographic separation of SWS cones into distinct dorsal and ventral fields was also obvious in retinal explants from newborn mice. However, the MWS opsin, identified by polyclonal antibody JH492, was expressed only in vitro when dorsal explants were harvested from P3 or older pups. Despite the absence of MWS opsin expression in newborn retinal cultures, there was no evidence of an increase in the numbers of SWS cones. To test if local diffusable cues could induce immature cones to express an aberrant opsin, dorsal and ventral retinal explants at different stages of maturation were cocultured during the incubation period. Neither the emergence of the cone fields nor the difference in the regional and temporal development of the MWS and SWS opsins was affected in these experiments. These results suggest that positional information in the retina and the opsin identity of cones is determined prior to birth and argue against the hypothesis that postnatal cones can be induced to express an aberrant opsin.

Increment threshold and purity discrimination spectral sensitivities of X-chromosome-linked color-defective observers

The goal of the study was to evaluate spectral opponency in nine X-chromosome-linked color-defective observers. The tasks included increment threshold spectral sensitivity on an achromatic background, heterochromatic flicker photometry, and colorimetric purity discrimination. With a task of heterochromatic flicker photometry, the anomalous trichromatic observers showed spectral sensitivity of the corresponding dichromat. The increment threshold spectral sensitivity and colorimetric purity discrimination data were analyzed using the concept of standard cone photopigment spectral sensitivities for normal and defective vision, and a model that postulates one cone-additive and two cone-antagonistic systems. The model incorporated a shift of the peak spectral sensitivity of the long-wavelength-sensitive (LWS) pigment (for protan observers) or of the middle-wavelength-sensitive (MWS) pigment (for deutan observers). Two dichromats and two anomalous trichromats did not show clear evidence of LWS vs MWS cone antagonism. Five anomalous trichromats showed such cone antagonism. Molecular genetic analysis of the opsin genes is presented for eight of the observers.

Mechanisms subserving temporal modulation sensitivity in silent-cone substitution.

Temporal contrast sensitivity data were collected with sine-wave-modulated lights for achromatic, chromatic, and silent-cone-substitution stimuli. Achromatic (556- and 642-nm lights in phase) and chromatic (556- and 642-nm lights in counterphase) modulation sensitivities were measured at a constant time-average retinal illuminance of 1256 trolands (Td) and chromaticity of 595 nm. These data were considered to represent isolated temporal responses of luminance and red-green chromatic channels, respectively. Silent cone substitution was achieved with counterphase modulation of the 556- and the 642-nm lights and by suitable adjustment of the modulations or the radiances of the two lights. (1) The peak modulation depth of the 642-nm light was reduced to silence the long-wavelength-sensitive (LWS) cone, and the peak modulation depth of the 556-nm light was reduced to silence the middle-wavelength-sensitive (MWS) cone. These protocols maintained the time-average retinal illuminance and chromaticity as for the control conditions. (2) The luminance of the 642-nm light was decreased to silence the LWS cone and was increased to silence the MWS cone. In this procedure the time-average retinal illuminance and chromaticity differ for the silenced-LWS-cone (1047 Td and 589.5 nm) and the silenced-MWS-cone (4358 Td and 622 nm) conditions. The response modulation of the achromatic and the chromatic channels was calculated for the silent-substitution conditions. The chromatic channel is more sensitive at low frequencies, with a transition to greater achromatic channel sensitivity near 13 Hz for the silenced-LWS-cone condition and near 6 Hz for the silenced-MWS-cone condition.(ABSTRACT TRUNCATED AT 250 WORDS)

Foveal cone detection statistics in color-normals and dichromats

We measured for six male observers, the psychometric functions for the detection of two simultaneously presented points of light. The test stimuli were two 1 min point sources separated by 17 min arc, and pulsed for 0.5 msec. The stimuli varied in wavelength from 500 to 620 nm. The psychometric functions were fit with a model that assumes ideal detection and the following properties: (1) Poisson-distributed quantal absorptions; (2) binomial sampling of foveal long-wavelength-sensitive (LWS) and middle-wavelength-sensitive (MWS) cones; (3) independent responses of the LWS and MWS cones; and (4) the Smith-Pokorny fundamentals for cone spectral sensitivies. Based on χ 2 fits to the psychometric functions for detecting neither, one or both of the two-point stimuli presented, estimates were derived for the minimum quantal catch by a single cone for detection ( C), the number of effective cones illuminated by a point stimulus at threshold ( N) and the proportion of central foveal cones of the LWS type ( P L ). Three observers were color-normal, two were protanopes and one was a deuteranope. A second deuteranope was included in the design but his data were too unreliable for an unambigous solution. The estimated quantal requirement C was consistently near 5 (4–6), and the effective number of illuminated cones always was 1 or 2. The plausible range of P L (98% confidence interval) for the color-normal observers was 0.48–0.68 (observer YY), 0.76–0.90 (observer MW) and 0.77–0.95 (observer DF). The best fitting P L solution for these observers were 0.61, 0.82 and 0.88, respectively. These were comparable to the values obtained from flicker photometric data. The best P L value for each of the protanopes was 0.00 and for the deuteranope the best P L value was 0.98.

Contribution of the retinal ON channels to scotopic and photopic spectral sensitivity

Visual information encoded by the middle-wavelength-sensitive (MWS) and long-wavelength-sensitive (LWS) cones in the primate retina are processed by both depolarizing (ON) and hyperpolarizing (OFF) bipolar cells. In contrast, signals from the short-wavelength-sensitive (SWS) cones and dark-adapted rod photoreceptors are thought to be carried almost exclusively by ON bipolar cells (Gouras & Evers, 1985). Consequently, it would be expected that functional inactivation of the retinal ON channels at the bipolar cell level would produce selective deficits in visual functions mediated by rods and SWS cones. We have examined this hypothesis by injecting rhesus monkeys with 2-amino-4-phosphonobutyric acid (APB), a pharmacological agent that reduces the responsiveness of retinal ON neurons, and psychophysically measuring the changes in spectral sensitivities. Under adaptation conditions that isolated rod function, APB caused, as expected, a substantial loss in rod-mediated spectral sensitivity. However, under photopic conditions, cone-mediated spectral sensitivity, including that associated with the SWS cones, was relatively unaffected. These results demonstrate distinct organizational differences between the rod and cone systems; specifically, they indicate that the rod system is more dependent upon retinal ON channels than the cone system. Our failure to find a selective visual deficit related to SWS cone function under photopic viewing conditions suggests that the OFF system can mediate stimulus detection throughout the visible spectrum and that the ability of the OFF system to process signals from the SWS cones has been underestimated.

Classification of complete and incomplete autosomal recessive achromatopsia.

We studied color vision in 32 patients with autosomal recessive achromatopsia. Color matching revealed complete achromatopsia (rod monochromasy) in ten patients (Group I) and incomplete achromatopsia in the remaining twenty-two patients. Amongst the incomplete achromats, were three groups distinguishable by their color matching. Patients in Group II were dichromats; their color matches were mediated by rods and MWS (middle-wavelength sensitive) cones. Patients in Groups III and IV were trichromats. Color matches of patients in Group III were mediated by rods, LWS (long-wavelength sensitive) cones and MWS cones. Group III patients showed no evidence of SWS (short-wavelength sensitive) cones. Color matches of patients in Group IV were mediated by rods, LWS cones and SWS cones; color matching did not reveal MWS cones.

New observations concerning red–green color defects

AbstractThe majority of research on X‐chromosomal defects has been with a 2° field of view. The studies concur in a view of dichromacy as a loss or reduction system, and anomalous trichromacy as an alteration system. According to this view, protanopes have no functional LWS (long‐wave‐length‐sensitive) photopigment; deuteranopes have no functional MWS (middle‐wavelength‐sensitive) photopigment. With larger fields of view, dichromats show trichromacy which is mediated by rods at low photopic luminances, and by anomalous cones for retinal illuminances above 1000 trolands. These studies suggest that the same (or similar) anomalous (L'WS) photopigment is formed at the protan locus and the same (or similar) anomalous (M'WS) photopigment is formed at the deutan locus for both the anomalous trichromatic and dichromatic forms of the defects. Family studies reveal that intrafamilial variability in chromatic discrimination is less than interfamilial variation. A possible interpretation of such studies is that multiple, rather than three or four, alleles may be involved in X‐chromosomal color‐vision defects, with the alleles determining the proportion of functional anomalous cones.