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PubMed1987

Ontogeny of the area centralis in the cat.

Previous studies have shown that after embryonic day 50 (E50) the kitten retina undergoes a relatively rapid period of maturation that begins at the area centralis and spreads toward the retinal periphery. The mechanism controlling this pattern is unknown. One possibility is that the area centralis is the most developmentally advanced region of retina from the earliest stages of eyecup formation. The present study tests this hypothesis. Retinae from ten kittens aged between E21 and E54 were sectioned horizontally at 10 micron and stained with haematoxylin and eosin. During the earliest stages of development in other vertebrate species, the oldest part of the retina is at the posterior pole of the eyecup, just dorsal to the optic fissure. At E21 in the kitten retina, this site appeared to be developmentally advanced by several criteria. A developmentally advanced region (DAR) was recognized at each subsequent age. It gradually became displaced temporally toward the site of the area centralis in the adult. Developmental events beginning here include the formation of cell-free channels at the vitreal margin; the appearance of the first ganglion cells and their axons; the formation of continuous inner and outer plexiform layers; the differentiation of cells in the ganglion cell, inner, and outer nuclear layers; and the cessation of cytogenesis at the outer limiting membrane. By E54 the DAR could be recognized as the area centralis by the criteria used in the adult. Between E25 and E37 cell density in the ganglion cell layer is higher in the DAR than at other retinal eccentricities. During the E40s cell density becomes uniform across the retina because more cells migrate into the ganglion cell layer in peripheral retina than in the DAR. Several observations suggest that cells added to the ganglion cell layer from E37 onward are not ganglion cells. The proportion of retinal area occupied by the DAR decreases geometrically between E21 and E54, indicating that this region grows at a slower rate than the rest of the retina. It is suggested that this differential growth combined with differential cell addition may be the principal mechanism responsible for generating centroperipheral cell gradients in the retina.

Animals↗

PubMed2001

Localization of neurotransmitters and calcium binding proteins to neurons of salamander and mudpuppy retinas.

We wished to identify the different types of retinal neurons on the basis of their content of neuroactive substances in both larval tiger salamander and mudpuppy retinas, favored species for electrophysiological investigation. Sections and wholemounts of retinas were labeled by immunocytochemical methods to demonstrate three calcium binding protein species and the common neurotransmitters, glycine, GABA and acetylcholine. Double immunostained sections and single labeled wholemount retinas were examined by confocal microscopy. Immunostaining patterns appeared to be the same in salamander and mudpuppy. Double and single cones, horizontal cells, some amacrine cells and ganglion cells were strongly calbindin-immunoreactive (IR). Calbindin-IR horizontal cells colocalized GABA. Many bipolar cells, horizontal cells, some amacrine cells and ganglion cells were strongly calretinin-IR. One type of horizontal cell and an infrequently occurring amacrine cell were parvalbumin-IR. Acetylcholine as visualized by ChAT-immunoreactivity was seen in a mirror-symmetric pair of amacrine cells that colocalized GABA and glycine. Glycine and GABA colocalized with calretinin, calbindin and occasionally with parvalbumin in amacrine cells.

Acetylcholine↗

PubMedDate not supplied

Light adaptation and color opponency of horizontal cells in the turtle retina.

Chromaticity-type (C-type) horizontal cells of the turtle retina receive antagonistic inputs from cones of different spectral types, and therefore their response to background illumination is expected to reflect light adaptation of the cones and the interactions between their antagonistic inputs. Our goal was to study the behavior of C-type horizontal cells during background illumination and to evaluate the role of wavelength in background adaptation. The photoresponses of C-type horizontal cells were recorded intracellularly in the everted eyecup preparation of the turtle Mauremys caspica during chromatic background illuminations. The voltage range of operation was either reduced or augmented, depending upon the wavelengths of the background and of the light stimuli, while the sensitivity to light was decreased by any background. The response-intensity curves were shifted to brighter intensities and became steeper as the background lights were made brighter regardless of wavelength. Comparing the effects of cone iso-luminant backgrounds on the Red/Green C-type horizontal cells indicated that background desensitization in these cells could not solely reflect background adaptation of cones but also depend upon response compression/expansion and changes in synaptic transmission. This leads to wavelength dependency of background adaptation in C-type horizontal cells, that is expressed as increased light sensitivity (smaller threshold elevation) and improved suprathreshold contrast detection when the wavelengths of the background and light stimuli were chosen to exert opponent effects on membrane potential.

Adaptation, Ocular↗

PubMed2000

Interindividual and topographical variation of L:M cone ratios in monkey retinas.

We analyzed the ratio of L:M cone photopigment mRNA in the retinas of Old World monkeys, using the method of rapid polymerase chain reaction-single-strand conformation polymorphism. The L:M cone pigment mRNA ratio in whole retina ranged from 0.6 to 7.0, with a mean of approximately 1.6 (standard deviation, +/- 0.56; n = 26). There was no change in this ratio with eccentricity up to 9 mm (approximately 45 degrees), though the ratio was approximately 30% greater in temporal than in nasal retina. The mRNA ratios are in good agreement with the L:M cone ratio in these same retinas, inferred from electrophysiological recordings of cone signal gain in horizontal cell interneurons. The correlation between mRNA ratios and physiological cone gain ratio supports the conclusion that both measures reflect the relative number of L and M cones.

Animals↗

PubMedDate not supplied

Turtle C-type horizontal cells act as push-pull devices.

Chromaticity (C-type) horizontal cells in the retina of cold-blooded vertebrates receive antagonistic inputs from cone photoreceptors of different spectral types leading to color opponency. The relative contribution of each spectral type of cones can be selectively altered by chromatic background illumination. Therefore, the spectral properties of C-type horizontal cells are expected to change when the intensity and color of ambient illumination are altered. In this study, we investigated the effects of chromatic background lights upon color opponency in Red/Green (RGH) and Yellow/Blue (YBH) C-type horizontal cells in the everted eyecup preparation of the turtle Mauremys caspica. Photoresponses were elicited by long-wavelength and short-wavelength light stimuli in the dark-adapted state and under conditions of chromatic background illumination. We found that the total voltage range. within which graded depolarizing and the hyperpolarizing photoresponses could be elicited, either increased or decreased depending upon the color of the background light. However, the maximal and minimal potential levels determined respectively by long-wavelength and short-wavelength light stimuli of supersaturating intensity remained unchanged, regardless of the wavelength and intensity of the background. These findings indicate that turtle C-type horizontal cells operate as push-pull devices. A sufficiently bright short-wavelength stimulus can push them all the way to the maximal hyperpolarizing level while a very bright long-wavelength stimulus can pull them towards the most depolarizing potential.

Adaptation, Ocular↗

PubMed1990

Calbindin and calretinin localization in retina from different species.

Calbindin-D28K and calretinin are homologous calcium-binding proteins localized in many neurons of the central nervous systems. We have compared polyclonal antibodies against calbindin and calretinin and have shown by western blots using purified calbindin and calretinin from rat that (1) anti-calretinin does not recognize calbindin and (2) anti-calbindin presents some cross-reactivity with calretinin. In this report, we have compared by immunohistochemistry the localization of both calcium-binding proteins in the retina of monkey, pig, sheep, rat, cat, pigeon, and salamander. These results are compared with previous data for chick. There are many differences between species and not within species, but some aspects of the distribution are conserved. All species, except rat and monkey, have some cones which contain calbindin only. Most species also have some bipolar cells containing calbindin only. Calretinin is rarely seen in photoreceptors or bipolar cells. All species have horizontal cells which contain calretinin or calbindin or both. All species have amacrine cells and ganglion cells containing one or other protein. In the cat ganglion cell layer, the calretinin antisera define a new, asymmetric, type of cell.

Animals↗

PubMed2001

The synaptic architecture of AMPA receptors at the cone pedicle of the primate retina.

Cone pedicles, the output synapses of cone photoreceptors, transfer the light signal onto the dendrites of bipolar and horizontal cells. Cone pedicles contain between 20 and 45 ribbon synapses (triads) which are the release sites for glutamate, the cone transmitter. Several hundred postsynaptic dendrites contact individual cone pedicles, and we studied the glutamate receptors expressed and clustered at these contacts, particularly the AMPA receptor subunits. Using immunocytochemistry and confocal imaging we were able to resolve individual triads within the cone pedicles by light microscopy. We studied their differences in L/M- and S-cones, and we counted the number of triads per pedicle across the retina. The presynaptic matrix protein bassoon, the synapse-associated membrane protein P84, and peanut agglutinin were used to specifically label synaptic ribbons, invaginating dendrites of horizontal cells and invaginating dendrites of ON-cone bipolar cells, respectively. Pre- and post-embedding immunocytochemistry and electron microscopy were used to localize the AMPA receptor subunits at the cone pedicle base. They were aggregated at three different postsynaptic sites: at horizontal cell invaginating contacts, at bipolar cell flat contacts, and at desmosome-like junctions underneath the cone pedicles. We also performed double-labeling experiments with the triad-specific markers and the antibodies against the AMPA receptor subunits. AMPA receptors were preferentially expressed by horizontal cells, and to a lesser extent by OFF-cone bipolar cells. We did not observe any cone-selective expression of AMPA receptor subunits postsynaptic to L/M- or S-cones, suggesting AMPA receptors are not the key to understanding trichromatic signaling in the primate retina.

Animals↗

PubMed1996

Histochemistry of NADPH-diaphorase--a marker for neuronal nitric oxide synthase--in the carp retina.

The nicotinamide adenine dinucleotide phosphate (NADPH) diaphorase histochemical technique was used as a marker to assess the distribution of nitric oxide synthase activity in the carp retina. NADPH-diaphorase activity was found to be present in photoreceptors (rods and cones), horizontal cells, amacrine cells, bipolar cells, Müller cells and ganglion cells. Staining was most prominent in the photoreceptor ellipsoids but was not confined to any particular subtype. The density of the staining within the inner plexiform layer (IPL) was determined by image analysis. There was a broad peak of activity in each sublamina of the IPL, but sublamina b appeared to be relatively more heavily stained. The results taken together suggest that the nitric oxide signalling system could have a broader involvement in retinal function than previously thought. Furthermore, nitric oxide may have a novel mode of action in the retina whereby it could be effective on cells (photoreceptors) that also synthesize it.

Animals↗

PubMed1989

Catecholamine-sensitive adenylate cyclase in the white perch (Roccus americanus) retina: evidence for beta-adrenergic and dopamine receptors linked to adenylate cyclase.

We have examined the catecholamine-sensitive adenylate cyclase in the retina of the white perch (Roccus americanus). Both dopamine and the beta-adrenergic agonist isoproterenol stimulate cyclic AMP accumulation in this retina, but serotonin, an indoleamine, and phenylephrine, an alpha-adrenergic agonist, had no effect. The stimulation of adenylate cyclase by isoproterenol is more potent and effective than that of dopamine. The effects of dopamine and isoproterenol are mediated via independent dopamine and beta-adrenergic receptors. Haloperidol, a dopamine antagonist, blocks the stimulatory effect of dopamine but not of isoproterenol. Conversely, propranolol, a beta-adrenergic antagonist, blocks the stimulatory effect of isoproterenol but not of dopamine. The effects of dopamine and isoproterenol are not additive. In fractions of purified horizontal cells we found evidence for dopamine receptors linked to adenylate cyclase but did not find evidence for the presence of cyclase coupled beta-adrenergic receptors. The cellular location of the beta-adrenergic receptors is unknown. Our findings demonstrate the existence of both beta-adrenergic and dopamine receptors coupled to adenylate cyclase in the white perch retina. However, we did not find either epinephrine or norepinephrine, endogenous ligands of the beta-receptor, to be present in retinal extracts subjected to HPLC.

Adenylyl Cyclases↗

PubMed1978

Perimetry of contrast detection thresholds of moving spatial sine wave patterns. IV. The influence of the mean retinal illuminance.

Contrast detection thresholds for moving sine wave gratings were obtained at the fovea and at eccentricities of 6 degrees, 21 degrees, and 50 degrees on the nasal horizontal meridian. The targets subtended from 30 X 30 minutes of arc up to 16 degrees X 16 degrees. Mean retinal illuminance was varied between 10 and 0.01 trolands. The transition from the de Vries-Rose to the Weber region occurs in the far peripheral visual field at a 2-3 decades lower illuminance level than at the fovea. The spatio-temporal contrast detection thresholds become comparable over the whole visual field if the mean distance between retinal ganglion cells is taken as a yardstick, and field width, spatial frequency, and quantum density are scaled accordingly. This means that at scotopic illuminance levels coarse or medium gratings are preferentially detected at other than foveal locations. (The fine gratings cannot be resolved at all at such levels.) It is argued that both electrophysiological and psychophysical evidence indicates that Weber behavior starts whenever some small fixed number of quantum absorptions occur within an area of 1 mean interganglion cell distance across. Or, equivalently, if a fixed small number of "neural quanta" enters a 100 X 100 micron2 area of the visual cortex.

Adult↗

PubMedDate not supplied

Early identification of retinal subtypes in the developing, pre-laminated chick retina using the transcription factors Prox1, Lim1, Ap2alpha, Pax6, Isl1, Isl2, Lim3 and Chx10.

In this study, antibodies toward the transcription factors Prox1, Lim1, Ap2alpha, Pax6, Isl1, Isl2, Lim3 and Chx10 were used to identify and distinguish between developing cell types in the pre-laminated chick retina. The spatio-temporal expression patterns were analysed from embryonic day 3 (E3) to E9, thus covering a time-span from the onset of retinal cell-fate determination to when retinal laminas can be distinguished. Most transcription factors were found at early stages of development, enabling us to trace various precursor cell populations throughout the lamination process. With time, each transcription factor expression became restricted to distinct laminas or sub-laminas of the maturing retina. These early emerging patterns were compared and found to be consistent with those of the hatched chick retina, where the outer nuclear layer label for Lim3, Isl1 and Isl2. In the inner nuclear layer, horizontal cells labeled for Prox1, Lim1, Isl1, Ap2alpha and Pax6, bipolar cell labeled for Lim3, Isl1 and Chx10 and amacrine cells labeled for Ap2alpha, Isl1 and Pax6. The ganglion cell layer labeled for Isl1, Pax6 and Isl2. The immunolabeling patterns of Lim3 and Isl2 have not previously been described in detail.

Animals↗

PubMed1985

Cholecystokinin in the cat retina. Action of exogenous CCK8 and localization of cholecystokinin-like immunoreactivity.

The effects of applying cholecystokinin (CCK8) iontophoretically onto cat retinal ganglion cells were studied in the optically intact eye of the cat. CCK8 suppressed both the maintained and the light evoked discharge of brisk ganglion cells, irrespective of their being on- or off-center, brisk-transient, or brisk-sustained and independent of the state of light adaptation. The inhibitory action of CCK8 in the cat retina is opposite from its excitatory action in other parts of the brain. Using an antiserum to cholecystokinin, immunoreactivity was localized in horizontal cells and amacrine cells of the cat retina. Inconsistently immunoreactivity also was found in ganglion cells and fibers.

Animals↗

PubMed1981

Gap junctions in the outer plexiform layer of te chick retina: thin section and freeze-fracture studies.

Previous studies have established that gap junctions between presumptive retinal neurons of the chick retina disappear during the course of embryogenesis. The present study examines the 2-3-week-old chick retina to determine if gap junctions are present in the outer plexiform layer of the more mature animal as would be in accordance with evidence from morphological and physiological studies on a variety of other vertebrates. Thin section and freeze-fracture techniques are used in a complementary manner to demonstrate that gap junctions are present between horizontal cell processes in the distal regions of the outer plexiform layer. These junctions appear to be between axon terminals and between spines that project from axon terminals to rods and double cones. Gap junctions are also observed between photoreceptors. They are seen on the synaptic terminals of all classes of cones and are located between the cone synaptic terminals and cone basal processes. Gap junctions are also seen between unidentified photoreceptor basal processes within the neuropil of both distal and proximal parts of the outer plexiform layer. Gap junctions are also present between cone synaptic terminals and deeply invaginated, vesicle-containing processes the origin of which remains to be determined.

Animals↗

PubMed1988

The role of the posterior vermis of monkey cerebellum in smooth-pursuit eye movement control. I. Eye and head movement-related activity.

1. The observation of smooth-pursuit eye and retinal image velocity signals in lobules VI and VII of the vermis has given rise to the hypothesis that a neural correlate of a target velocity signal exists in this region of the cerebellum (29). However, activity signaling head velocity is also required to regenerate a target velocity signal. Vermal Purkinje cell activity was, therefore, recorded during the performance of paradigms designed to dissociate head movement-related responses. 2. The activity of 107 Purkinje cells was found to be related to horizontal head velocity. Of these, 52% increased their discharge rate for ipsilaterally directed passive head movement (type I), and 48% were excited by contralateral head movements (type II). 3. In five Purkinje cells in which sufficient data were obtained, cell discharge rate increased monotonically with head velocity over the range of 5-40 deg/s. The sensitivity to head velocity at 0.4 Hz +/- 25 deg/s averaged approximately 0.5 spikes.s-1/deg.s-1 in a larger sample of cells (n = 39). The sensitivities to head velocity, at this same frequency and velocity, of type I and type II Purkinje cells were comparable at 0.44 and 0.51 spikes.s-1/deg.s-1, respectively. 4. The Purkinje cell responses led head velocity by an average of 12 degrees at 0.4 Hz +/- 25 deg/s of passive head rotation. The phase shifts associated with type I and II responses were similar with phase leads of 13 and 9 degrees with respect to head velocity, respectively. 5. A linear interaction of smooth-pursuit eye and head velocity signals was observed during the performance of a variety of antiphase and inphase eye and head movement paradigms. The results support the conclusion that some Purkinje cells in lobules VI and VII of the cerebellar vermis encode a gaze velocity signal. Contributions of the head velocity signal to the regeneration of target velocity are considered in a companion paper (32).

Animals↗

PubMed1989

Organized arrangement of orientation-sensitive relay cells in the cat's dorsal lateral geniculate nucleus.

We studied the physiological orientation biases of over 700 relay cells in the cat's dorsal lateral geniculate nucleus (LGNd). Relay cells were sampled at regular intervals along horizontally as well as vertically oriented electrode penetrations in a fashion analogous to that used previously in studies of visual cortex (Hubel and Wiesel, 1962). The strengths of the orientation biases and the distributions of the preferred orientations were determined for different classes of relay cells, relay cells in different layers of the LGNd, and relay cells subserving different parts of the visual field. We find that, at the population level, LGNd cells exhibit about the same degree of orientation bias as do the retinal ganglion cells providing their inputs (see also Soodak et al., 1987). Also, as in the retina (Levick and Thibos, 1982; Leventhal and Schall, 1983), most LGNd cells tend to prefer stimuli oriented radially, i.e., parallel to the line connecting their receptive fields to the area centralis projection. However, the radial bias in the LGNd is weaker than in the retina. Moreover, there is a relative overrepresentation of cells preferring tangentially oriented stimuli in the LGNd but not in the retina. As a result of the overrepresentation of cells preferring radial and tangential stimuli, the overall distribution of preferred orientations varies in regions of the LGNd subserving different parts of the visual field. Reconstructions of our electrode penetrations provide evidence that, unlike in the retina, cells having similar preferred orientations are clustered in the LGNd. This clustering is apparent for all cell types and in all parts of laminae A and A1. The tendency to cluster according to preferred orientation is evident for cells preferring radially, intermediately, and tangentially oriented stimuli and thus is not simply a reflection of the radial bias evident among retinal ganglion cells at the population level. It is already known that cells having inputs from different eyes, on-center, off-center, X-, Y-, W-type, and color-sensitive ganglion cells are distributed nonrandomly in the LGNd of cats and monkeys (for review, see Rodieck, 1979; Stone et al., 1979; Lennie, 1981; Stone, 1983). The finding that relay cells having similar preferred orientations are also distributed nonrandomly suggests that the initial sorting of virtually all properties segregated in visual cortex may begin in the LGNd.

Animals↗

PubMed1984

Retinal ganglion cells that project to the dorsal lateral geniculate nucleus in the macaque monkey.

Horseradish peroxidase was deposited in the optic nerve to retrogradely label and reveal the dendritic form of all classes of ganglion cell, or it was injected into the dorsal lateral geniculate nucleus to reveal only those classes projecting to the thalamus. The results were compared with those of the accompanying paper in which the ganglion cells projecting to the midbrain are selectively revealed. Two major classes of ganglion cells are described, the P alpha and P beta cells. For both classes dendritic field size increases with eccentricity from the fovea and there is no overlap in the two classes at any given eccentricity. Cell body size shows a similar mean difference but with a slight overlap. Both cell bodies and dendritic fields are larger along the temporal horizontal meridian than the nasal horizontal meridian, for P alpha and for P beta cells, but these differences are reduced when naso-temporal differences in ganglion cell density are taken into account, that is, size correlates closely with density. Injections restricted to the parvocellular layers of the lateral geniculate nucleus labelled almost exclusively P beta cells, whereas injections confined to the magnocellular layers labelled almost exclusively P alpha cells. As midbrain injections label no P beta cells and few P alpha cells it can be shown that about 80% of ganglion cells are P beta cells projecting to parvocellular lateral geniculate nucleus, and that about 10% are P alpha cells projecting to magnocellular layers. The coverage factor, that is the number of cells covering each point on the retina, varied from 1.9-2.3 for P beta cells, and from 2-7 for P alpha cells. Comparing the results with those of comparable investigations on cats and rabbits shows a much clearer segregation of the terminal targets of different classes of ganglion cell in monkeys, the greatest difference being the absence in the monkey of a projection to the geniculate from gamma- and epsilon-like cells. Further, axons which branch and innervate both thalamus and midbrain are rare in monkeys but common in other mammals. Comparing the results with those from physiological investigations suggests that the P beta cells correspond to colour-opponent cells, whereas P alpha cells correspond to the achromatic broad-band magnocellular cells.

Animals↗

PubMed1984

Development and degeneration of retina in rds mutant mice: electron microscopy.

In the retina of mice homozygous for the retinal degeneration slow (rds) gene, receptor outer segments failed to develop and typical disc structures were never observed. However, cilia surrounded by a plasma membrane were regularly present. At the time when outer segments grew in the normal retina, the optic ventricle in the mutant showed an accumulation of membrane-bound vesicles of varying size and density. The vesicles declined in frequency at later stages of degeneration. After initial growth, the inner segments in the mutant retina remained stunted but maintained their morphological differentiation, including the ciliary structures. Their number declined with loss of visual cells. In the mutant retina development of synapses, between the receptor cells and the processes of the horizontal cells and the bipolar cell dendrites, followed in normal sequence. Formation of spherule terminals of the rods, with one triad synapse, and of pedicle terminals of the cones, with multiple triad synapses, was recorded in the same way as in the normal retina. With loss of visual cells in the mutant retina, some of the profiles of the surviving spherule terminals showed an increased occurrence of two, three, or more synaptic sites. This growth resulted from enlargement and branching of the ribbons and sprouting from the postsynaptic elements. Similar changes were not observed in the pedicle terminals. The pigment epithelial cells in the mutant mice appeared initially normal, but phagolysosomal structures were absent. However, various inclusion bodies appeared within the pigment epithelial cells following degenerative changes in the retina. In some places, the basal infoldings deepened their furrows while thinning the cytoplasmic part of the epithelium. Failure to form the outer segments and the subsequent lysis of the visual cells appear to be the primary effects of the rds gene, whereas the synaptic growth in the sperule terminals and the changes in the pigment epithelium appear to be consequential to those defects.

Animals↗

PubMed1975

[Reactions of frog retinal rods to light].

The amplitude and shape of the rod photoresponses were investigated at different intensity, duration, wavelength and diameter of the light spot. Intracellular recordings were performed from the green and red rod outer sigments. An increase in the light intensity produced an increase in the amplitude and a decrease in the time rise of the response. An intensive light flash suppressed for a time the rod sensitivity to test flashes. At light spots of large diameter (100--1500 mum) delayed depolarizing deflection was observed. The latter was conditioned by illumination of receptor surroundings and, apparently, was due to horizontal cells activity. The possible role of the depolarizing effect of the rod surroundings illumination is discussed.

Animals↗