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PubMed1979

The emergence, localization and maturation of neurotransmitter systems during development of the retina in Xenopus laevis. I. Gamma aminobutyric acid.

The high-affinity uptake, biosynthesis and release of GABA have been studied in the retina of Xenopus laevis. In the mature retina, [3H]-GABA is accumulated predominantly by horizontal cells. A second population of cells located in the inner nuclear layer (possibly a type of amacrine cell) also showed a specific GABA uptake. In addition, this retina contains significant activities of L-glutamic acid decarboxylase and also releases [3H]-GABA in response to increasing K+ concentrations in the medium. We have followed the appearance and maturation of these GABA-ergic properties during embryonic development of this retina. Our results indicate that these properties emerge in a precise temporal pattern during retinal differentiation: the specific neuronal uptake of GABA precedes GABA synthesis which is followed by K+-stimulated GABA release.

Animals↗

PubMed1994

Subcellular localization of GABAA receptor on bipolar cells in macaque and human retina.

The subcellular distribution of GABAA receptor in the macaque and human retina was studied by immunocytochemistry with monoclonal antibodies for the alpha and beta subunits with a particular focus on bipolar cells. Immunoreactivity to GABAA receptor was present on dendritic tips of all bipolar cells. The stain was strongest on bipolar membranes in apposition to horizontal cell processes. Stain was concentrated on the tips of flat and invaginating cone bipolar cells at the base of the cone pedicle and on the invaginating tips of rod bipolar cells. Stain on the cone pedicle membrane was restricted to sites of apposition to stained bipolar dendrites; pedicle membrane in apposition to horizontal cell processes was unstained. Stain was also present on bipolar axon terminals in both on and off strata of the inner plexiform layer. All bipolar cell somas stained faintly; horizontal and Müller cell somas were unstained. The alpha and beta subunits distributed similarly in monkey and human retina. Presence of GABAA receptor on the bipolar dendritic tips suggests that horizontal cells directly affect bipolar cells. Thus, GABAA receptor might mediate the receptive field surround of both off and on bipolar cells. Presence of GABAA receptor on bipolar axon terminals suggests that much of the inhibition feeding back from GABAergic amacrine to bipolar cells is GABAA-mediated.

Animals↗

PubMed1989

Control of the optokinetic reflex by the nucleus of the optic tract in primates.

Physiological and anatomical experiments clearly established the existence of a pretectal relay of visual information to the ipsilateral inferior olive in the macaque monkey. Horseradish peroxidase injected into the inferior olivary nucleus retrogradely labelled neurons in the nucleus of the optic tract (NOT) and the dorsal terminal nucleus of the accessory optic tract (DTN). The response characteristics of NOT-DTN neurones are described in this chapter. The visual receptive fields of neurones in NOT and DTN in anaesthetized and paralysed macaque monkeys prefer horizontal ipsiversive movements of single objects or whole field random dot patterns, i.e. neurones in the left NOT-DTN prefer leftward movements and vice versa. The directional tuning widths of NOT-DTN neurones are very broad. Directions withing a mean range of 127 +/- 25 degrees visual angle elicit response strengths greater than 50% of the maximal response. Visual latencies to reversals in directions of stimulus movement are in a range from 40 to 80 ms (mean 61 +/- 13 ms). Combining two visual stimuli by moving a random dot pattern and a single bar of light simultaneously but in opposite directions causes NOT-DTN neurones to respond to each stimulus as soon as it moves in the cell's preferred direction. The reduced overall response strengths indicate additional inhibitory interactions. All NOT-DTN neurones can be activated from each eye. Interactions between the two eyes are modest and unspecific. Optical speeds of stimulus movement vary for different NOT-DTN neurones (4-60 deg/s). The effective range of speeds is broad (0.1-400 deg/s for the total population). With oscillating horizontal stimulation NOT-DTN neurones follow repetition rates up to 4 Hz. Receptive fields are mostly large (20-40 degrees visual angle), include the fovea, and extend up to 20 degrees into the ipsilateral hemifield. The sensitivity to moving stimuli is highest near the fovea. Our results thus indicate that direction selective cells in the NOT and DTN have all the properties and connections which are necessary and sufficient to control the stability of the image on the retina by supplying retinal slip information to the velocity storage integrator in the brainstem (Raphan et al., 1979).

Animals↗

PubMed1998

Amino acid signatures in the normal cat retina.

PURPOSE: To establish a nomogram of amino acid signatures in normal neurons, glia, and retinal pigment epithelium (RPE) of the cat retina, guided by the premise that micromolecular signatures reflect cellular identity and metabolic integrity. The long-range objective was to provide techniques to detect subtle aberrations in cellular metabolism engendered by model interventions such as focal retinal detachment. METHODS: High-performance immunochemical mapping, image registration, and quantitative pattern recognition were combined to analyze the amino acid contents of virtually all cell types in serial 200-nm sections of normal cat retina. RESULTS: The cellular cohorts of the cat retina formed 14 separable biochemical theme classes. The photoreceptor --> bipolar cell --> ganglion cell pathway was composed of six classes, each possessing a characteristic glutamate signature. Amacrine cells could be grouped into two glycine- and three gamma-aminobutyric acid (GABA)-dominated populations. Horizontal cells possessed a distinctive GABA-rich signature completely separate from that of amacrine cells. A stable taurine-glutamine signature defined Müller cells, and a broad-spectrum aspartate-glutamate-taurine-glutamine signature was present in the normal RPE. CONCLUSIONS: In this study, basic micromolecular signatures were established for cat retina, and multiple metabolic subtypes were identified for each neurochemical class. It was shown that virtually all neuronal space can be accounted for by cells bearing characteristic glutamate, GABA, or glycine signatures. The resultant signature matrix constitutes a nomogram for assessing cellular responses to experimental challenges in disease models.

Alanine↗

PubMedDate not supplied

Characterization of the rod photoresponse isolated from the dark-adapted primate ERG.

The a-wave of the human dark-adapted ERG is thought to derive from activity of rod photoreceptors. However, other sources within the retina could potentially perturb this simple equation. We investigated the extent to which the short-latency dark-adapted rod a-wave of the primate ERG is dominated by the rod photoresponse and the applicability of the phototransduction model to fit the rod a-wave. Dark-adapted Ganzfeld ERGs were elicited over a 5-log-unit intensity range using short bright xenon flashes, and the light-adapted cone responses were subtracted to isolate the rod ERG a-wave. Intravitreal 4-phosphono-butyric acid (APB) and cis-2,3-piperidine-dicarboxylic acid (PDA) were applied to isolate the photoreceptor response. The Hood and Birch version of the phototransduction model, Rmax[1 - e(-I x S x (t-t(eff)))2], was fitted to the a-wave data while allowing Rmax and S to vary. Three principle observations were made: (1) At flash intensities > or =0.77 log sc-td-s the leading edge of the normalized rod ERG a-wave tracks the isolated photoreceptor response across the first 20 ms or up to the point of b-wave intrusion. The rod ERG a-wave was essentially identical to the isolated receptor response for all intensities that produce peak responses within 14 ms after the flash. (2) The best fit of sensitivity (S) was not affected by APB and/or PDA, suggesting that the inner retina contributes very little to the dark-adapted a-wave. (3) APB always reduced the maximum dark-adapted a-wave amplitude (by 15-30%), and PDA always increased it (by 7-15%). Using the phototransduction model, both events can be interpreted as a scaling of the photoreceptor dark current. This suggests that activity of postreceptor cells somehow influences the rod dark current, possibly by feedback through horizontal cells (although currently not demonstrated for the rod system), or by altering the ionic concentrations near the photoreceptors, or by neuromodulator effects mediated by dopamine or melatonin.

Aminobutyrates↗

PubMed1989

Role of cerebellar flocculus in adaptive interaction between optokinetic eye movement response and vestibulo-ocular reflex in pigmented rabbits.

Sustained sinusoidal oscillation of a striped cylindrical screen around a stationary, alert pigmented rabbit with certain parameters (for 4 h. 5 degrees, 7.5 degrees, 10 degrees peak-to-peak, 0.1 or 0.2 Hz) adaptively modified not only the horizontal optokinetic reflex (HOKR) but also the horizontal vestibuloocular reflex (HVOR). The major effects thus obtained during 4 h were an increase in the HOKR gain by 0.23, and that of the HVOR gain by 0.18. Bilateral destruction of floccular Purkinje cells with microinjection of kainic acid abolished these effects on both HOKR and HVOR. Single unit activities of floccular Purkinje cells were recorded from the floccular areas related to horizontal eye movements (H-zone) with local stimulus effects. Most H-zone Purkinje cells normally exhibited modulation of simple spike discharge in phase with screen velocity and out of phase with turntable velocity. Sustained screen oscillation (7.5 degrees, 0.1 Hz) for 1 h increased the simple spike responses not only to screen but also to turntable oscillation. No such changes were observed in other floccular areas. These observations suggest that sustained optokinetic stimulations induce adaptation of HVOR through an interaction of retinal slip and head velocity signals within the flocculus or its related neuronal tissues.

Action Potentials↗

PubMed1980

Fine structure of the compound eyes and interfacetal mechanoreceptors of Cicindela tranquebarica Herbst (Coleoptera: Cicindelidae).

The structure of the compound eyes of adult Cicindela tranquebarica Herbst was examined by use of light, scanning, and transmission electron microscopy. Each ommatidium of these photopic eyes is eucone and has a "subcorneal layer" situated between the corneal lens and crystalline cone. A distal rhabdomere consisting only of microvilli from retinula cell seven, a more proximal, rectangular, fused rhabdom formed from six retinula cells, and a basal, eighth retinula cell with a spherical rhabdomere comprise the light sensitive portions of the ommatidium. The "subcorneal layer" consists of lamellae of endocuticular microfibrils and, in surface view, shows 11 concave polygons. Proximal extensions of the crystalline thread form inter-retinular fibres containing microtubules between retinula cells 1/2, 3/4, 5/6, and 7/1. The primary pigment cells are devoid of pigment granules, but are rich in rough endoplasmic reticulum. Proximal to each retinula cell nucleus are two basal bodies, one perpendicular to the other. The more proximal basal body extends two fibrillar feet proximally which fuse to form a horizontally-banded ciliary rootlet extending the length of the retinula peripheral to the rhabdom. Each ommatidium is surrounded by 16 secondary pigment cells. Interfacetal mechanoreceptors between some adjacent lenses each have a single bipolar neuron, with a dendritic sheath, tubular body, cilium, outer and inner sheath cells, and an axon surrounded by a neurilemma sheath cell.

Animals↗

PubMed1991

4B2: a monoclonal antibody to ganglion and bipolar cells in the retina of the cat, generated by intrasplenic immunization.

Using tissue from the inner layers of the retina as the immunogen, we have prepared a monoclonal antibody which is selective for two classes of neuron in the cat retina. The antibody, termed 4B2, was generated by intrasplenic injection, demonstrating that neuron-specific antibodies can be elicited by this technique, which requires only micrograms of immunogen. The 4B2 binds to the somas, dendrites and axons of ganglion cells. Double labelling with 4B2 and with a retrograde tracer injected into the retino-recipient nuclei of the brain demonstrates that, of the cell classes in the ganglion cell layer, 4B2 labels only ganglion cells, and that, amongst ganglion cells, 4B2 is strongly selective for large (alpha-) and medium-sized (beta- and perhaps gamma-) cells. Double labelling with 4B2 and with markers for amacrine cells confirms tht 4B2 does not label amacrine cells. Double labelling with 4B2 and with anti-GFAP confirms that 4B2 does not label the macroglia of the retina. The 4B2 also labels bipolar cells, showing their somas, dendrites and axons. The axons of the labelled cells terminate in large boutons, in the innermost part of the inner plexiform layer and in the ganglion cell layer. This level of termination suggests that 4B2 labels rod bipolars, and perhaps a subgroup of cone bipolars. Double labelling with 4B2 and with markers for amacrine, horizontal and Müller cells indicates that other cell classes with somas in the inner nuclear layer remain unlabelled. In addition, we have examined the ontogeny of 4B2 labelling in the cat retina. The developmental sequence of labelling with 4B2 follows the sequence in which the cells are born, first ganglion and then bipolar cells.

Animals↗

PubMed1990

Orientation bias of neurons in the lateral geniculate nucleus of macaque monkeys.

The purpose of this investigation was to analyze the influence of stimulus orientation on the responses of individual neurons in the monkey's lateral geniculate nucleus (LGN). Our specific goals were to assess the prevalence and the degree of orientation tuning in the monkey LGN and to determine if the preferred stimulus orientations of LGN neurons varied as a function of receptive-field position. The primary motivation for this research was to gain insight into the receptive-field configuration of LGN neurons and consequently into the neural mechanisms which determine the spatial organization of LGN receptive fields in primates. In both the parvocellular and magnocellular layers, the responses of the majority of individual neurons to sine-wave gratings varied as a function of stimulus orientation. The influence of stimulus orientation was, however, highly dependent on the spatial characteristics of the stimulus; the greatest degree of orientation bias was observed for spatial frequencies higher than the cell's optimal spatial frequency. On a population basis, the degree of orientation bias was similar for all major classes of LGN neurons (e.g. ON vs. OFF center; parvocellular vs. magnocellular) and did not vary systematically with receptive-field eccentricity. At a given receptive-field location, LGN neurons, particularly cells in the parvocellular laminae, tended to prefer either radially oriented stimuli or stimuli oriented more horizontally than their polar axis. Our analyses of the orientation-dependent changes in spatial-frequency response functions, which was based on the Soodak et al., (1987; Soodak, 1986) two-dimensional, difference-of-Gaussian receptive-field model, suggested that the orientation bias in LGN neurons was due to an elongation of the receptive-field center mechanism which in some cases appeared to consist of multiple subunits. Direct comparisons of the orientation-tuning characteristics of LGN cells and their retinal inputs (S potentials) indicated that the orientation bias in the monkey LGN reflects primarily the functional properties of individual retinal ganglion cells. We conclude that orientation sensitivity is a significant property of subcortical neurons in the primate's geniculo-cortical pathway.

Action Potentials↗

PubMedDate not supplied

Localization of AMPA-selective glutamate receptor subunits in the cat retina: a light- and electron-microscopic study.

The distribution of AMPA-selective glutamate receptor subunits was studied in the cat retina using antisera against GluR1 and GluR2/3. Both antisera were localized in postsynaptic sites in the outer plexiform layer (OPL) as well as the inner plexiform layer (IPL). Immunoreactivity for GluR1 was seen in a subpopulation of OFF cone bipolar cells and a number of amacrine and ganglion cells. Within the IPL, processes staining for GluR1 received input from OFF and ON cone bipolar cells but not from rod bipolars. Labeling for GluR2/3 was seen in horizontal cells, an occasional cone bipolar cell, and numerous amacrine and ganglion cells. In the IPL, GluR2/3 staining was postsynaptic to cone bipolar cells in both sublaminae. AII amacrine cells which receive rod bipolar input were also labeled for GluR2/3. With both antisera, staining was limited to a single member of the bipolar dyad complex, providing morphological evidence for functional diversity in glutamatergic pathways.

Animals↗

PubMed1986

Retinal neurochemistry of three elasmobranch species: an immunohistochemical approach.

We surveyed retinas of Raja erinacea, Mustelus canis, and Squalus acanthias for neurotransmitter substances by using antisera directed against the substances themselves or against their synthesizing enzymes. Both the peroxidase-antiperoxidase (PAP) and indirect fluorescent techniques were employed to visualize the primary antisera. In all three species positive results were obtained with antisera directed against tyrosine hydroxylase (TOH), glutamic acid decarboxylase (GAD), serotonin (5-HT), and leucine enkephalin (Lenk). Antisera directed against glucagon, neurotensin, beta-endorphin, vasoactive intestinal peptide, or bombesin failed to show any specific staining. Immunoreactivity was located in amacrine, interplexiform, and horizontal cells as well as in axons of the optic fiber layer. The four antisera labelled different amacrine cell classes, distinguished on the bases of perikaryal morphology and the distribution of cell processes in the inner plexiform layer (IPL). Amacrine cells that labelled with the same marker were seen to have different morphologies in the species studied. Thus, TOH-like immunoreactivity was distributed in layers 1, 3, and 5 of the IPL in Mustelus but only in layers 1 and 3 in Raja retina. GAD-like immunoreactivity was found diffusely over all layers of the IPL in Raja, but in Mustelus it was confined primarily to layers 1, 3, and 5 of the IPL. Lenk- and 5-HT-like immunoreactivities showed similar species variations. Two neurochemical classes of interplexiform cell were identified in this study. In Mustelus GAD-like and Lenk-like immunoreactive interplexiform cells were seen whereas in Raja only GAD-positive interplexiform cells were detected. In squalus no unequivocal demonstration of any interplexiform cell was made with these antisera. The GAD antiserum also labelled a subset of the horizontal cells in the dorsal retina of Raja. TOH and 5-HT-antisera labelled axons in the optic fiber layer of all three species but reactive ganglion cell perikarya were not identified.

Animals↗

PubMed1980

A layered model for visual processing in avian retina.

Visual processing in avian retina is interpreted by means of a layered model in which: a) outer layers provide with spatio temporal fast and retarded versions of the stimuli incident on the retina; a possibility is that horizontal cells are involved in isotropically generating the retarded version which is transversally translated; b) prominent specialization of ganglion cells is the result of local non-linear lateral interaction at the inner plexiform layer, mediated by amacrines which return, also isotropically, the translated retarded signals. Small though systematic deviations in the sites of the lateral interaction result in anisotropic but uniform receptive fields for some ganglion cells. A simple though general expression for the model is derived which includes the various types of recorded avian ganglion retinal cells responses, which also permits a unified interpretation of visual processing in avian and cat's retinae.

Animals↗

PubMed2002

Cone neurite sprouting: an early onset abnormality of the cone photoreceptors in the retinal degeneration mouse.

PURPOSE: Mutations in many rod genes can cause inherited blinding neurodegeneration in the retina characterized by sequential death of rod and cone photoreceptors. This study was to examine the morphological changes of the cone photoreceptors in retinal degeneration (rd1) mice caused by rod-specific cGMP phosphodiesterase beta-subunit gene mutation and to gain insights into the early cellular events underlying the secondary cone death. METHODS: Transgenic mice that have their living cones labeled by the green fluorescent protein (GFP) transgene and carry the homozygous rd1 mutation were generated, and identified by PCR analysis of the mouse tail DNA and PCR coupled Dde I digestion. The morphology of cone cells in live and fixed retinas from developing and adult mice was examined with fluorescence and scanning laser confocal microscopy. Some fixed mouse retinas were also examined by immunocytochemical staining. Volume images from the confocal three-dimensional (3D) data sets were processed with IMARIS software for 3D view of the detailed cone cell morphology. RESULTS: The cone photoreceptors in the rd1 retinas exhibited a novel process of neurite sprouting, in addition to the general pathological changes of cone degeneration such as shortening and loss of cone outer and inner segments, and loss and death of the cones. The cones gave rise to prominent neurite outgrowth from their axons and synaptic pedicles as well. Most neurites had beaded varicosities along their length and some terminated as bulbous structures. Some cone pedicles showed abnormally elongating and branching processes. The degenerating cones were disorganized, and migrated into the inner nuclear layer. Some cone neurites extended horizontally and appeared to contact the rod bipolar cells, while others projected into the inner plexiform layer. The aberrant cone sprouting started from P8 when rod degeneration generally began, and became evident by P10. In contrast, this abnormal cone neurite sprouting was not observed in the examined control mice that did not carry the rd1 mutation. Double-labeling with cone cell-specific peanut agglutinin confirmed that the fluorescent cells expressing the GFP in the rd1 retinas were indeed the cone photoreceptors. CONCLUSIONS: Cone photoreceptors in the rd1 mice underwent a remarkable process of neurite sprouting that appeared to start before the onset of cone cell death and persisted throughout the course of cone degeneration. This novel process of cone neurite sprouting may be a part of the early cellular events leading to the cone photoreceptor death in retinal degeneration of the rd1 mice.

3',5'-Cyclic-GMP Phosphodiesterases↗

PubMed1995

The appearance of rod opsin during monkey retinal development.

PURPOSE: To determine the temporal and spatial pattern of rod opsin appearance in Macaca monkey retina. METHODS: Frozen sections from fetal day (Fd) 55 to adulthood (birth = Fd168) containing the entire horizontal meridian were stained using Rho4D2 monoclonal antiserum visualized with immunofluorescent labeling. At Fd66, Fd79, and Fd89, retinal samples taken at known eccentricities were studied from the opposite eye using standard electron microscope methods. RESULTS: Rod opsin was detected at Fd66 in or near the fovea, and a second focus appeared at Fd75 to Fd77 near the optic disc in the nasal rod ring. The earliest opsin appeared in the apical stubs, which resembled the apical connecting cilium in the electron microscope. Staining of the entire cell body membrane, including the synaptic spherule, was present 4 to 7 days later. Opsin expression had a nasal bias with rods at the nasal ora labeled at Fd140, whereas temporal ora was not labeled until Fd155. Cell body labeling disappeared by Fd132 across central retina but persisted into the first postnatal year in far peripheral retina. Outer segment (OS) length measurements showed that rods in the rod ring had the longest OS between Fd115 and postnatal week 9. Rod OS at all retinal eccentricites continued to elongate between 11 months of age and adulthood. CONCLUSIONS: Rod opsin expression follows a foveal-to-peripheral gradient beginning at Fd66 and ending near birth. Rod opsin is detected first in the connecting cilium and slightly later in the entire cell membrane, and then cell membrane labeling disappears as the heavily labeled OS elongates. Although the first OS appear on rods near the fovea, these OS still are short at birth and do not reach adult length until after 2 years of age. The longest OS at birth are found on rods at the rod ring, suggesting that this region could have higher scotopic sensitivity than central retina at birth.

Aging↗

PubMed1997

Localization of D1 dopamine receptors in the chicken retina.

PURPOSE: The localization of dopamine D1 receptors (DIR) in the chicken retina was examined using an anti-human DIR monoclonal antibody and PAP techniques. RESULTS: A clear band of staining was seen in the outer plexiform layer, as well as cellular staining in the outer-most part of the inner nuclear layer, probably in a subset of horizontal cells. Many different amacrine cell bodies were labelled in the inner one-third of the inner nuclear layer. There was also extensive staining in the inner plexiform layer, which showed some striation. Occasional labelled ganglion cells were also detected. CONCLUSION: Localization of D1-dopamine receptors has been shown in the chicken retina.

Animals↗

PubMedDate not supplied

Multiple neuronal connexins in the mammalian retina.

Gap junctions are abundant in the mammalian retina and many neuronal types form neural networks. Several different neuronal connexins have now been identified in the mammalian retina. Cx36 supports coupling in the AII amacrine cell network and is essential for processing rod signals. Cx36 is probably also responsible for photoreceptor coupling. Horizontal cells appear to be extensively coupled by either Cx50 or Cx57. These results indicate that multiple neuronal connexins are expressed in the mammalian retina and that different cell types express different connexins.

Amacrine Cells↗

PubMed1982

Ratfish retina--intracellular recordings and HRP injections in an isolated, superfused all-rod retina.

Hyperpolarizing responses to light were studied by intracellular recording in the isolated, superfused retina of the ratfish (Hydrolagus colliei). Two of these hyperpolarizing units were identified as horizontal cells by iontophoretic injection of horseradish peroxidase (HRP). These cells had rather large cell bodies (15 x 30 micrometer), elliptical dendritic arborizations measuring 150 x 300 micrometers and no axons. Since their physiological receptive fields were found to be at least 2.15 mm in diameter, it appears likely that either the photoreceptors or the horizontal cells are electrically coupled. Electron microscopy of HRP-injected horizontal cells showed their dendrites to end laterally in ribbon synapses of rods only and revealed dendro-dendritic contacts resembling gap junctions between injected and uninjected horizontal cells. The spectral sensitivity function of a dark-adapted horizontal cell can be described by a Dartnall nomogram based on a retinene pigment with lambda max = 473 nm. These findings are consistent with the histological observation that the ratfish retina appears to contain only rod photoreceptors.

Animals↗

PubMed1974

Control of retinal sensitivity. II. Lateral interactions at the outer plexi form layer.

Test stimuli, presented at the center of the bipolar cell receptive field, spanning less than 2 log units of intensity, elicit the full range of graded response. The intensity range of test stimuli that elicits the graded response depends upon the background conditions. A higher range of log test intensities is required to elicit the graded bipolar response in the presence of surround backgrounds. But surround backgrounds can also serve to unsaturate the bipolar response and thereby increase sensitivity under certain conditions. The results suggest that a second stage of sensitivity-control is mediated by the horizontal cell system at the outer plexiform layer, concatenated with the effects of adaptation in the photoreceptors.

Adaptation, Physiological↗