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Gerald H Jacobs

Publications and source records attributed to Gerald H Jacobs.

16 recordsLinked to original sources

Visual pigments of marine carnivores: pinnipeds, polar bear, and sea otter.

Rod and cone visual pigments of 11 marine carnivores were evaluated. Rod, middle/long-wavelength sensitive (M/L) cone, and short-wavelength sensitive (S) cone opsin (if present) sequences were obtained from retinal mRNA. Spectral sensitivity was inferred through evaluation of known spectral tuning residues. The rod pigments of all but one of the pinnipeds were similar to those of the sea otter, polar bear, and most other terrestrial carnivores with spectral peak sensitivities (lambda(max)) of 499 or 501 nm. Similarly, the M/L cone pigments of the pinnipeds, polar bear, and otter had inferred lambda(max) of 545 to 560 nm. Only the rod opsin sequence of the elephant seal had sensitivity characteristic of adaptation for vision in the marine environment, with an inferred lambda(max) of 487 nm. No evidence of S cones was found for any of the pinnipeds. The polar bear and otter had S cones with inferred lambda(max) of approximately 440 nm. Flicker-photometric ERG was additionally used to examine the in situ sensitivities of three species of pinniped. Despite the use of conditions previously shown to evoke cone responses in other mammals, no cone responses could be elicited from any of these pinnipeds. Rod photoreceptor responses for all three species were as predicted by the genetic data.

Amino Acid Sequence↗

Polymorphic New World monkeys with more than three M/L cone types.

Most New World (platyrrhine) monkeys have M/L cone photopigment polymorphisms that map directly into individual variations in visual sensitivity and color vision. We used electroretinogram flicker photometry to examine M/L cone photopigments in the New World monkey Callicebus moloch (the dusky Titi). Like other New World monkeys, this species has an M/L cone photopigment polymorphism that reflects the presence of X-chromosome opsin gene alleles. However, unlike other platyrrhines in which three M/L photopigments are typical, Callicebus has a total of five M/L cone photopigments. The peak sensitivity values for these pigments extend across the range from 530 to 562 nm. The result is an enhanced array of potential color vision phenotypes in this species.

Animals↗

Photoreceptors and photopigments in a subterranean rodent, the pocket gopher (Thomomys bottae).

Pocket gophers (Thomomys bottae) are rodents that spend much of their lives in near-lightless subterranean burrows. The visual adaptations associated with this extreme environment were investigated by making anatomical observations of retinal organization and by recording retinal responses to photic stimulation. The size of the eye is within the normal range for rodents, the lens transmits light well down into the ultraviolet, and the retina conforms to the normal mammalian plan. Electroretinogram recording revealed the presence of three types of photopigments, a rod pigment with a spectral peak of about 495 nm and two types of cone pigment with respective peak values of about 367 nm (UV) and 505 nm (medium-wavelength sensitive). Both in terms of responsivity to lights varying in temporal frequency and in response recovery following intense light adaptation, the cone responses of the pocket gopher are similar to those of other rodents. Labeling experiments indicate an abundance of cones that reach densities in excess of 30,000 mm-2. Cones containing UV opsin are found throughout the retina, but those containing medium-wavelength sensitive opsin are mostly restricted to the dorsal retina where coexpression of the two photopigments is apparently the rule. Rod densities are lower than those typical for nocturnal mammals.

Adaptation, Ocular↗

Influence of cone pigment coexpression on spectral sensitivity and color vision in the mouse.

The mouse retina contains both middle-wavelength-sensitive (M) and ultraviolet-sensitive (UV) photopigments that are coexpressed in cones. To examine some potential visual consequences of cone pigment coexpression, spectral sensitivity functions were measured in mice (Mus musculus) using both the flicker electroretinogram (ERG) and behavioral discrimination tests. Discrimination tests were also employed to search for the presence of color vision in the mouse. Spectral sensitivity functions for the mouse obtained from ERG measurements and from psychophysical tests each reveal contributions from two classes of cone having peak sensitivities (lambda(max)) of approximately 360 and 509-512 nm. The relative contributions of the two pigment types to spectral sensitivity differ significantly in the two types of measurements with a relationship reversed from that often seen in mammals. Mice were capable of discriminating between some pairs of spectral stimuli under test conditions where luminance-related cues were irrelevant. Since mice can make dichromatic color discriminations, their visual systems must be able to exploit differences in the spectral absorption properties among the cones. Complete selective segregation of opsins into individual photoreceptors is apparently not a prerequisite for color vision.

Animals↗

Evolution of vertebrate colour vision.

Recent years have witnessed a growing interest in learning how colour vision has evolved. This trend has been fuelled by an enhanced understanding of the nature and extent of colour vision among contemporary species, by a deeper understanding of the paleontological record and by the application of new tools from molecular biology. This review provides an assessment of the progress in understanding the evolution of vertebrate colour vision. In so doing, we offer accounts of the evolution of three classes of mechanism important for colour vision--photopigment opsins, oil droplets and retinal organisation--and then examine details of how colour vision has evolved among mammals and, more specifically, among primates.

Animals↗

Genetically engineered mice with an additional class of cone photoreceptors: implications for the evolution of color vision.

Among eutherian mammals, only primates possess trichromatic color vision. In Old World primates, trichromacy was made possible by a visual pigment gene duplication. In most New World primates, trichromacy is based on polymorphic variation in a single X-linked gene that produces, by random X inactivation, a patchy mosaic of spectrally distinct cone photoreceptors in heterozygous females. In the present work, we have modeled the latter strategy in a nonprimate by replacing the X-linked mouse green pigment gene with one encoding the human red pigment. In the mouse retina, the human red pigment seems to function normally, and heterozygous female mice express the human red and mouse green pigments at levels that vary between animals. Multielectrode array recordings from heterozygous female retinas reveal significant variation in the chromatic sensitivities of retinal ganglion cells. The data are consistent with a model in which these retinal ganglion cells draw their inputs indiscriminately from a coarse-grained mosaic of red and green cones. These observations support the ideas that (i) chromatic signals could arise from stochastic variation in inputs drawn nonselectively from red and green cones and (ii) tissue mosaicism due to X chromosome inactivation could be one mechanism for driving the evolution of CNS diversity.

Action Potentials↗

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.

Animals↗

Cone pigment variations in four genera of new world monkeys.

Previous research revealed significant individual variations in opsin genes and cone photopigments in several species of platyrrhine (New World) monkeys and showed that these in turn can yield significant variations in color vision. To extend the understanding of the nature of color vision in New World monkeys, electroretinogram flicker photometry was used to obtain spectral sensitivity measurements from representatives of four platyrrhine genera (Cebus, Leontopithecus, Saguinus, Pithecia). Animals from each genus were found to be polymorphic for middle to long-wavelength (M/L) sensitive cones. The presence of a short-wavelength sensitive photopigment was established as well so these animals conform to the earlier pattern in predicting that all male monkeys are dichromats while, depending on their opsin gene array, individual females can be either dichromatic or trichromatic. Across subjects a total of five different M/L cone pigments were inferred with a subset of three of these present in each species.

Adaptation, Ocular↗

Topography of photoreceptors and retinal ganglion cells in the spotted hyena (Crocuta crocuta).

The spatial distributions of photoreceptors and retinal ganglion cells were examined in the spotted hyena (Crocuta crocuta). Two populations of cones were identified by immunocytochemical labeling. The hyena retina contains approximately 2.3 million middle- to long-wavelength sensitive (M/L) cones that reach peak densities of about 7,500/mm(2) in the vicinity of the optic nerve head. A sparser population of short-wavelength sensitive (S) cones, totaling about 0.3 million, was also detected. There is a striking disparity in the spatial distributions of the two cone types with S cones achieving peak density in a region located well below the optic nerve head. The differences in the spatial distributions of the two cone types have implications both for visual sensitivity and for color vision. Hyena rods outnumber cones by about 100:1 with rod density falling off modestly along a central-peripheral gradient. Ganglion cells were identified in retinal wholemounts by Nissl staining patterns. Their distribution defines a prominent visual streak with highest spatial packing (approx. 4,200/mm(2)) in an area centralis that is located in the temporal retina. The total number of ganglion cells is estimated at about 260,000. Using standard assumptions the maximum spatial resolution of the spotted hyena is calculated to be about 8.4 cycles/degree, a value similar to estimates obtained for other terrestrial carnivores.

Animals↗

Cone photoreceptor recovery after experimental detachment and reattachment: an immunocytochemical, morphological, and electrophysiological study.

PURPOSE: To compare the morphologic and functional recovery of the retina after detachment and reattachment in an animal with a cone-dominant retina, the ground squirrel. METHODS: Ground squirrel (Spermophilus beecheyi) retinas were detached for 1 day and reattached for 7, 35, or 96 days (n = 2, each time point). Flicker ERGs were recorded 1 day after the detachment and at various times after reattachment. Contrast-response functions were measured for isochromatic modulation and for selective modulation of short-wavelength-sensitive (S) and middle-wavelength-sensitive (M) cones. At the end of the experiment, retinas were prepared for light microscopy or immunocytochemical staining with antibodies to rod opsin, S and M cone opsins, cytochrome oxidase, synaptophysin, glial fibrillary acidic protein (GFAP), cellular retinaldehyde-binding protein (CRALBP), interphotoreceptor-binding protein (IRBP), and peanut agglutinin lectin (PNA). Photoreceptor density maps were created from wholemount preparations labeled with biotinylated PNA and anti-S cone opsin. Cell counts of photoreceptor nuclei and cone outer segments (OS) were compared with flicker ERG data. Cell death was examined by the TUNEL method. RESULTS: Reattachment stopped photoreceptor cell death and reversed the disruption of interphotoreceptor matrix as well as the redistribution of Müller cell proteins. It also activated some astrocytes based on anti-GFAP staining. S- and M-cone OS showed a gradual recovery in length after reattachment, and this recovery continued to the longest time points examined. ERG contrast gains also recovered after reattachment, but these reached asymptotic levels by approximately a week after reattachment. There were significant correlations between outer nuclear layer (ONL) cell counts and ERG contrast gains. No differences were noted in the indices of recovery of M and S cones. CONCLUSIONS: The ERG can be used to follow specifically the changes in the retina that occur after retinal detachment and reattachment.

Animals↗

Opsin gene and photopigment polymorphism in a prosimian primate.

A recent genetic investigation found some species of prosimian to have an opsin gene polymorphism [Nature 402 (1999) 36]. In the present study the functional implications of this finding were explored in a correlated investigation of opsin genes and spectral sensitivity measurements of a diurnal prosimian, Coquerel's sifaka (Propithecus verreauxi coquereli). Spectra recorded using electroretinogram (ERG) flicker photometry reveal a cone photopigment polymorphism paralleling an opsin gene polymorphism detected by molecular methods. This species has two middle-to-long-wavelength cone pigments with peak sensitivities of about 545 and 558 nm and a short-wavelength-sensitive cone with a peak at about 430 nm. The distribution of these pigments among animals predicts the presence of both dichromatic and trichromatic forms of color vision.

Animals↗

An animal model for studying cone function in retinal detachment.

In people, retinal detachment often leads to a significant loss in cone-based vision. Most of the animal models commonly used for studying the consequences of retinal detachment have rod-dominated retinas. The purpose of this investigation was to evaluate the possibility that the ground squirrel, a rodent with a heavily cone-dominated retina, might provide a useful model for studying cone function in retinal detachment. Corneal ERGs were recorded from ground squirrels for large-field temporal modulations presented on a computer-controlled color monitor. Modulations were chosen to selectively stimulate either of the two classes of cone found in the ground squirrel retina. Under these test conditions, large and reliable cone ERGs could be readily recorded. In animals in which the retina had been surgically detached, the loss of cone signal was directly related to the number of cones in the detachment zone relative to the total cone population and that relationship did not differ for short-wavelength sensitive (S) and middle-wavelength sensitive (M) cones. Surgical reattachment produced a progressive recovery of cone-based signals. The ground squirrel seems likely to provide a useful animal model for studying the dynamics of cone function in retinal detachment and subsequent events.

Animals↗

Spectral properties and retinal distribution of ferret cones.

The spectral mechanisms of the ferret (Mustela putorious furo) were studied with electroretinogram (ERG) flicker photometry. Variations in adaptation state and flicker rate were used to define corneally based spectral sensitivities for the three classes of receptor present in the retina of this mustelid-rods (lambdamax = 505 nm), S cones (430 nm), and L cones (558 nm). The retinal distributions of the two classes of cone were determined using opsin antibody labeling. Ferret retinas contain a total of about 1.3 million cones with L cones outnumbering S cones in a ratio of approximately 14:1. ERGs were also recorded using 18.75-Hz flickering stimuli that were designed to isolate signals from individual cone classes. The contrast/response functions for signals originating from both S and L cones were linear over low-to-moderate levels of contrast, but with greatly different slopes for the two cone types. The L:S contrast gain ratio derived from a comparison of these slopes, as well as inferences drawn from another experiment in which responses to various combinations of L- and S-cone activation were analyzed, suggest that contributions of these two cone types to the flicker ERG have a relative weighting of about 4:1 to 5:1 (L/S).

Animals↗

Cone pigment polymorphism in New World monkeys: are all pigments created equal?

Most platyrrhine monkeys have a triallelic M/L opsin gene polymorphism that underlies significant individual variations in color vision. A survey of the frequencies of these polymorphic genes suggests that the three alleles occur with equal frequency among squirrel monkeys (subfamily Cebinae), but are not equally frequent in a number of species from the subfamily Callitrichinae. This departure from equal frequency in the Callitrichids should slightly increase the ratio of dichromats to trichromats in the population and significantly alter the relative representation of the three possible dichromatic and trichromatic phenotypes. A particular feature of the inequality is that it leads to a relative increase in the number of trichromats whose M/L pigments have the largest possible spectral separation. To assess whether these trichromatic phenotypes are equally well equipped to make relevant visual discriminations, psychophysical experiments were run on human observers. A technique involving the functional substitution of photopigments was used to simulate the discrimination between fruits among a background of leaves. The goal of the simulation was to reproduce in the cones of human observers excitations equivalent to those produced in monkey cones as the animals view fruit. Three different viewing conditions were examined involving variations in the relative luminances of fruit and leaves and the spectrum of the illuminant. In all cases, performance was best for simulated trichromacies including M/L pigments with the largest spectral separation. Thus, the inequality of opsin gene frequency in Callitrichid monkeys may reflect adaptive pressures.

Animals↗

Rod and cone function in coneless mice.

Transgenic coneless mice were initially developed to study retinal function in the absence of cones. In coneless mice created by expressing an attenuated diphtheria toxin under the control of flanking sequences from the human L-cone opsin gene, a small number of cones (3-5% of the normal complement) survive in a retina that otherwise appears structurally quite normal. These cones predominantly ( approximately 87% of the total) contain UV-sensitive photopigment. ERG recordings, photoreceptor labeling, and behavioral measurements were conducted on coneless and wild-type mice to better understand how the nature of this alteration in receptor complement impacts vision. Signals from the small residual population of UV cones are readily detected in the flicker ERG where they yield signal amplitudes at saturation that are roughly proportional to the number of surviving cones. Behavioral measurements show that rod-based vision in coneless mice does not differ significantly from that of wild-type mice, nor does their rod system show any evidence of age-related deterioration. Coneless mice are able to make accurate rod-based visual discriminations at light levels well in excess of those required to reach cone threshold in wild-type mice.

Animals↗

L and M cone proportions in polymorphic New World monkeys.

Platyrrhine monkeys typically have only a single X-chromosome opsin gene. Alleles of this gene code for multiple versions of middle- to long-wavelength cone photopigments. X-chromosome inactivation provides heterozygous females with a retinal mosaic of cones containing either of two types of M and L pigment, thus establishing the photopigment basis for trichromatic color vision. This study examined the proportions of L and M cones created by this process. For that purpose, electroretinogram flicker photometry was used to obtain complete spectral sensitivity functions from 60 heterozygous female monkeys drawn from seven genera of platyrrhine monkeys. To obtain estimates of cone proportions, these functions were subsequently fit with linear combinations of L and M cone fundamentals that were derived from similar recordings made on conspecific animals having only one type of M/L pigment. Consistent with a random X-chromosome inactivation process, the average L:M cone weighting across the sample was close to unity. At the same time, there were significant individual variations in L:M cone proportions. The genesis of this variation and its implications for seeing are discussed.

Animals↗