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Retinal ganglion cell shrinkage in glaucoma.

SUMMARY: There has been some debate concerning the selective loss of retinal ganglion cells belonging to the magnocellular pathway in early glaucoma. Although histologic studies of retinal ganglion cells in experimental and human glaucoma have suggested selective loss of the larger cells and, by implication, selective damage to the magnocellular pathway, this has not been confirmed using psychophysical tests. Recent studies of retinal ganglion cell morphology in experimental glaucoma provide evidence that retinal ganglion cells undergo morphologic changes prior to cell death; cell volume is reduced in surviving cells with corresponding reductions in the size of the axon and dendritic tree. The magnitude of these changes is consistent with cell shrinkage as an explanation for the apparent selective damaged reported in earlier studies. It is also likely that widespread changes in the retinal ganglion cell population precede cell death, which will affect the physiologic behavior of these cells.

Animals↗

Psychophysical investigation of ganglion cell loss in early glaucoma.

PURPOSE: To evaluate ganglion cell loss in early glaucoma using a variety of psychophysical tests and to identify optimal perimetric technique(s) for detection of early glaucomatous visual function loss. METHODS: Five perimetric tests, short wavelength automated perimetry (SWAP), temporal modulation perimetry (TMP), frequency doubling technology perimetry (FDT), detection acuity perimetry (DAP), and resolution acuity perimetry (RAP) were compared in their ability to discriminate between normal individuals and patients with early glaucoma or glaucoma suspects. Comparisons were also made by their ability to produce repeatable defects. The tests examined different visual functions that are likely to be mediated by different retinal ganglion cell subpopulations, thereby permitting examination of hypotheses of ganglion cell death in early glaucoma. RESULTS: All visual field tests demonstrated high performance in separating glaucoma patients from normal individuals. SWAP, TMP, FDT, and DAP provided the greatest discrimination between normal individuals and high- and low-risk glaucoma suspects. However, SWAP, TMP, and FDT obtained better consistency across the various analysis approaches (global indices and pointwise) than DAP and RAP. Of all the test types, FDT exhibited the highest proportion of repeatable abnormal test locations, with poor confirmation rates achieved by DAP and RAP. CONCLUSION: The performance of SWAP, FDT, and TMP suggests that these test types may all be suitable for detection of early loss of visual function in glaucoma. Ganglion cell subpopulations with lower levels of redundancy and/or those with larger cell sizes offer the most parsimonious explanation for earliest ganglion cell losses occurring in glaucoma.

Cell Death↗

Is word recognition correlated with the number of surviving spiral ganglion cells and electrode insertion depth in human subjects with cochlear implants?

OBJECTIVES/HYPOTHESIS: Speech perception scores using cochlear implants have ranged widely in all published series. The underlying determinants of success in word recognition are incompletely defined. Although it has been assumed that residual spiral ganglion cell population in the deaf ear may play a critical role, published data from temporal bone specimens from patients have not supported this hypothesis. The depth of insertion of a multichannel cochlear implant has also been suggested as a clinical variable that may be correlated with word recognition. In the current study these correlations were evaluated in 15 human subjects. STUDY DESIGN: Retrospective review of temporal bone histopathology. METHODS: Temporal bones were fixed and prepared for histological study by standard techniques. Specimens were then serially sectioned and reconstructed by two-dimensional methods. The spiral ganglion cells were counted, and the depth of insertion of the cochlear implant as measured from the round window was determined. Correlation analyses were then performed between the NU6 word scores and spiral ganglion cell counts and the depth of insertion. RESULTS: The segmental and total spiral ganglion cell counts were not significantly correlated (P > .50) with NU6 word scores for the 15 subjects. Statistically significant correlations were not achieved by separate analysis of implant types. Similarly, no significant correlation between the depth of insertion of the electrode array and postoperative NU6 word score was identified for the group. CONCLUSION: Although it is unlikely that the number of residual spiral ganglion cell counts is irrelevant to the determination of word recognition following cochlear implantation, there are, clearly, other clinical variables not yet identified that play an important role in determining success with cochlear implantation.

Aged↗

Transneuronal degeneration of beta retinal ganglion cells in the cat.

Transneuronal retrograde degeneration of retinal ganglion cells was investigated following neonatal visual cortex ablation in the cat. After a survival time of at least 18 months, retinal ganglion cells projecting to the thalamus were labelled by retrograde transport of horseradish peroxidase. Filled ganglion cells were classified into alpha, beta and gamma types on the basis of dendritic morphology. In normal cats, alpha cells made up 8-10% of the total population in the sample area, beta cells made up 64-67% and gamma cells made up 23-27%. In retinae of visual cortex-ablated cats, normal numbers of alpha and gamma cells were present, but the beta cell population was depleted by 90% of normal. Thalamic projections of surviving retinal ganglion cells were investigated by anterograde transport of tritiated proline injected into the eye. In these animals, ablation of visual cortex resulted in almost complete degeneration of laminae A and A1 of the dorsal lateral geniculate nucleus. In the radioautographic material, projections from the retina to the degenerated parts of laminae A and A1 were barely detectable. Survival of some ganglion cell populations and death of others after neonatal visual cortex ablation may be explained in terms of the pattern of projections of the different cell types. We conclude that the majority of beta cells degenerate following visual cortex ablation because of removal of cells in the dorsal lateral geniculate nucleus which form their sole or principal target. Alpha and gamma cells and 10% of beta-cells survive because of extensive collateral projections to targets other than cells of the laminae A and A1 of dorsal lateral geniculate nucleus.

Animals↗

Changes in goldfish retinal ganglion cells during axonal regeneration.

Recent work suggests that mammalian retinal ganglion cells may become more like developing ganglion cells in form while regenerating through a peripheral nerve graft. We have injected Lucifer Yellow into regenerating ganglion cells of goldfish to look for similar changes. Within three weeks of injury, we saw dye-coupling to nearby cells, which is a common developmental feature in many species. Dendrites and axons, which in most mature ganglion cells are smooth, became varicose and hairy, like those examined in mammalian development. Secondary axons arose later, not only as side-branches of the primary axon but also from the soma, as in mammalian development and regeneration. Since, in fish, these responses are clearly an intrinsic part of functional regeneration, their equivalence in fish and mammals strengthens the view that a similar regenerative competence may exist in the retinal ganglion cells of all vertebrates.

Amidines↗

Temporal and mosaic distribution of large ganglion cells in the retina of a daggertooth aulopiform deep-sea fish (Anotopterus pharao).

The daggertooth Anotopterus pharao (Aulopiformes: Anotopteridae) is a large, piscivorous predator that lives within the epipelagic zone at night. In this species, the distribution of retinal ganglion cells has been examined. An isodensity contour map of ganglion cells shows that the cells concentrate in a slightly ventral region of the temporal retina. The region of high ganglion cell density contains 4.07 x 10(3) cells mm(-2), and the resulting visual acuity is 3.5 cycles deg(-1). Outside the area centralis, conspicuously large ganglion cells (LGCs) are observed in the temporal margin of the retina. The LGCs are regularly arrayed, and displaced into the inner plexiform layer. Thick dendrites extend into the outer part (sublamina a) of the inner plexiform layer. In the retinal whole mount, the total number of LGCs is 1590 (90.7 cm specimen), and the mean size of the LGCs is about four times larger than that of the ordinary ganglion cells. The morphological appearance of the LGCs was similar to the off-type alpha cells of the cat retina. The function of these distinctive LGCs is discussed in relation to specific head-up feeding behaviour.

Animals↗

The rabbit rectococcygeus: a ganglion-free parasympathetically innervated preparation.

1 Isolated, desheathed preparations of the rabbit rectococcygeus muscle were relatively insensitive to spasmogens other than muscarinic drugs. Transmural stimulation with 1-50 pulses (0.2-0.4 ms at 10 Hz) elicited graded twitches which were abolished by tetrodotoxin and were therefore neurogenic; longer pulses sometimes triggered tetrodotoxin-resistant myogenic contractions.2 Twitches elicited by 0.2-0.4 ms pulses were due to post-ganglionic excitation because they were not reduced by hexamethonium, pentolinium or dimethyltubocurarine, or by ganglion-paralyzing concentrations of nicotine.3 The acetyl- and butyryl-cholinesterase activities of the rectococcygeus were determined manometrically and could be selectively inhibited by BW 284C51 (1:5-bis-(4-allyl-dimethylammonium-phenyl)-pentan-3-one dibromide) and iso-OMPA (tetramonoisopropylpyrophosphortetramide), respectively. Single-pulse twitches were greatly potentiated in amplitude and duration only when both cholinesterases were inhibited.4 The preparations could not be made to contract by nicotine (2.1-21 muM) even after cholinesterase inhibition, suggesting an absence of ganglion-cells; with nicotine (105-210 muM) small, atropine-susceptible responses were elicited, which were non-ganglionic because they were not reduced by tetrodotoxin.5 Rectococcygeus preparations that had been treated with physostigmine released acetylcholine into the bath fluid on electrical stimulation.6 The motor transmission was paralyzed by botulinum toxin (Type A) and abolished by atropine; the block of muscarinic receptors by atropine was effective against both endogenous and exogenous acetylcholine.7 Inhibitory adrenoceptors and scanty motor alpha-adrenoceptors were detected in the smooth muscle.8 Strong inhibitions of motor transmission and of rhythmic activity were produced by noradrenaline (but not by tyramine), by isoprenaline, and, after phentolamine, also by adrenaline and phenylephrine. These inhibitions were only slightly reduced by propranolol and rather more by pindolol.9 Experiments on preparations retaining their extrinsic nerve supply suggest an absence of ganglionic relays in the last 1-2 cm of the motor nerve pathway to this muscle.10 Some contrasting properties of the adjacent caudo-anal muscle, including the poor motor responses to transmural stimulation, are described.

Acetylcholine↗

Effect of ganglionic blocking compounds on in-vivo fluid secretion in the rat small intestine.

It is well-known that enteric, secreto-motor nerves mediate cholera toxin-induced fluid secretion in the rat small intestine. This notion is, in part, derived from experiments on anaesthetized animals in which the response to cholera toxin was antagonized by the ganglionic nicotinic receptor antagonist, hexamethonium. In the current study, such anti-secretory action of ganglionic blocking compounds was analysed in an experiment designed to minimize any possible negative effect of general anaesthesia on intestinal secretion. Rats were anaesthetized with ether for 5-10 min, during which time a jejunal loop (10-12 cm) was constructed. The loop was challenged with one of the secretagogues, cholera toxin, prostaglandin E1 (PGE1) or okadaic acid. Saline (control) or either of the ganglionic blockers, hexamethonium and chlorisondamine, was administered intravenously. The rats were killed 5 h (cholera toxin) or 1.5 h (PGE1 and okadaic acid) after challenge, and the amount of fluid accumulated in the loops was determined. Cholera toxin-induced secretion was unchanged by hexamethonium but reduced by approximately 80% by chlorisondamine. The difference in effect between the two blockers might relate to the duration of ganglionic blockade. Chlorisondamine blocked secretion induced by either PGE1 or okadaic acid by approximately 60%. It is suggested that the anti-secretory effect of ganglionic blocking compounds might be a result of blockade of secreto-motor nerves but other mechanisms, for example interference with haemodynamic factors, cannot be ruled out.

Alprostadil↗

Indoleamine-mediated reciprocal modulation of on-centre and off-centre ganglion cell activity in the retina of the cat.

The effects of applying the indoleamines serotonin (5-hydroxytryptamine) and 5-methoxy-N,N-dimethyltryptamine (MDT) ionophoretically onto cat retinal ganglion cells were studied in the optically intact eye of the cat. Serotonin suppressed both the spontaneous activity and light-evoked discharge of on-centre ganglion cells, regardless of the visual stimulus used; on the other hand, it enhanced the activity of off-centre ganglion cells. MDT produced the opposite effects, i.e. it enhanced the activity of on-centre ganglion cells but suppressed that of off-centre ganglion cells. Much of the effect of serotonin on light-evoked discharge can be attributed to an alteration of maintained discharge. There was no major difference in the way brisk-sustained (X) and brisk-transient (Y) cells responded to the application of serotonin and MDT, although the effects of the indoleamines on brisk-transient (Y) cells were generally weaker than on brisk-sustained (X) cells. In view of the fact that the action of serotonin is similar to the action of a gain control system, a possible contribution of indoleamine-accumulating neurones to gain control in the cat retina is discussed.

Action Potentials↗

Responses of macaque ganglion cells to movement of chromatic borders.

1. We have measured responses of macaque ganglion cells to moving borders under conditions designed to simulate the minimally distinct border (MDB) task. 2. Extending previous results, we show that minimization of responses of phasic ganglion cells of the magnocellular (MC)-pathway obey the photometric laws of transitivity and additivity. 3. To equal luminance borders, a residual response was present in MC-pathway cells analogous to the second harmonic response seen in these neurones with temporal chromatic modulation. It was proportional to the tritanopic purity difference (magnitude of delta Pt, the rectified middle- to long-wavelength cone opponent signal) between the two colours on either side of the border. For a delta Pt of one, the mean residual response was equivalent to the response evoked by achromatic borders of about 14% luminance contrast. Both these properties of the MC-pathway closely resemble psychophysical estimates as to the distinctness of equal luminance borders. 4. We show how MC-pathway cell responses could be used centrally to support the MDB task. It was difficult to generate a model from responses of tonic ganglion cells of the parvocellular (PC)-pathway which would support the task. 5. The MDB task is still possible psychophysically after blurring the retinal image. Although blurring the border spatially smeared the responses of MC-pathway ganglion cells and reduced their amplitude, responses still went through a minimum close to equal luminance. Thus, blurring the image did not affect the ability of MC-pathway cells to support the task. Blurring the retinal image decreased the 'sharpness' of the border response of tonic, PC-pathway ganglion cells, but response amplitude was unaffected. Response features indicative of centre-surround organization were attenuated. A central mechanism reliant on centre-surround field structure of PC-pathway cells would thus not be able to support the task after blurring. 6. Taken together, these results strongly suggest that the MC-pathway forms the sole physiological substrate of the MDB task, and any contribution of the PC-pathway is, indeed, minimal.

Animals↗

Hyperacuity in cat retinal ganglion cells.

Cat X retinal ganglion cells that can resolve sine gratings of only 2.5 cycles per degree can nevertheless respond reliably to displacements of a grating of approximately 1 minute of arc. This is a form of hyperacuity comparable in magnitude to that seen in human vision. A theoretical analysis of this form of hyperacuity reveals it to be a result of the high gain and low noise of ganglion cells. The hyperacuity expected for the best retinal ganglion cells is substantially better than that observed in behavioral experiments. Thus the brain, rather than improving on the retinal signal-to-noise ratio by pooling signals from many ganglion cells, is unable to make use of all the hyperacuity information present in single ganglion cell responses.

Animals↗

The effects of morphine on sympathetic transmission in the stellate ganglion of the cat.

The aim of this study was to investigate which of the processes involved in synaptic transmission are affected by morphine in concentrations comparable to those used during surgical procedures. The effects of morphine sulfate on ganglionic transmission were studied in the stellate ganglion of the cat using intracellular and extracellular recordings in vitro. The neurons of the stellate ganglion were depolarized using preganglionic nerve stimulation, postganglionic nerve stimulation, and intracellular stimulation before and after introduction of morphine sulfate (up to 20 micrograms/mL). Tissue concentrations of morphine were estimated using radiolabeled morphine. Axonal transmission and the excitability of the postganglionic neurons to direct intracellular stimulation was not affected at the concentrations of morphine studied. In addition, morphine had a dose-dependent depolarizing effect on the resting membrane potential of most of the neurons in the stellate ganglion. Such neuronal depolarizations alone could initially produce excitation in some cell populations, followed by inhibition, secondary to the membrane depolarization, leading to depression of sympathetic nerve activity. The overall ganglionic transmission as recorded using an evoked potential was biphasic. At low doses morphine facilitated transmission, while at larger doses morphine attenuated evoked potentials. These effects do not appear to be mediated through classical opiate receptors since they are not blocked by naloxone.

Animals↗

Spike-dependent GABA inputs to bipolar cell axon terminals contribute to lateral inhibition of retinal ganglion cells.

The inhibitory surround signal in retinal ganglion cells is usually attributed to lateral horizontal cell signaling in the outer plexiform layer (OPL). However, recent evidence suggests that lateral inhibition at the inner plexiform layer (IPL) also contributes to the ganglion cell receptive field surround. Although amacrine cell input to ganglion cells mediates a component of this lateral inhibition, it is not known if presynaptic inhibition to bipolar cell terminals also contributes to surround signaling. We investigated the role of presynaptic inhibition by recording from bipolar cells in the salamander retinal slice. TTX reduced light-evoked GABAergic inhibitory postsynaptic currents (IPSCs) in bipolar cells, indicating that presynaptic pathways mediate lateral inhibition in the IPL. Photoreceptor and bipolar cell synaptic transmission were unaffected by TTX, indicating that its main effect was in the IPL. To rule out indirect actions of TTX, we bypassed lateral signaling in the outer retina by either electrically stimulating bipolar cells or by puffing kainate (KA) directly onto amacrine cell processes lateral to the recorded cell. In bipolar and ganglion cells, TTX suppressed laterally evoked IPSCs, demonstrating that both pre- and postsynaptic lateral signaling in the IPL depended on action potentials. By contrast, locally evoked IPSCs in both cell types were only weakly suppressed by TTX, indicating that local inhibition was not as dependent on action potentials. Our results show a TTX-sensitive lateral inhibitory input to bipolar cell terminals, which acts in concert with direct lateral inhibition to give rise to the GABAergic surround in ganglion cells.

Action Potentials↗

Electrical stimulation of mammalian retinal ganglion cells with multielectrode arrays.

Existing epiretinal implants for the blind are designed to electrically stimulate large groups of surviving retinal neurons using a small number of electrodes with diameters of several hundred micrometers. To increase the spatial resolution of artificial sight, electrodes much smaller than those currently in use are desirable. In this study, we stimulated and recorded ganglion cells in isolated pieces of rat, guinea pig, and monkey retina. We used microfabricated hexagonal arrays of 61 platinum disk electrodes with diameters between 6 and 25 microm, spaced 60 microm apart. Charge-balanced current pulses evoked one or two spikes at latencies as short as 0.2 ms, and typically only one or a few recorded ganglion cells were stimulated. Application of several synaptic blockers did not abolish the evoked responses, implying direct activation of ganglion cells. Threshold charge densities were typically <0.1 mC/cm2 for a pulse duration of 100 micros, corresponding to charge thresholds of <100 pC. Stimulation remained effective after several hours and at high frequencies. To show that closely spaced electrodes can elicit independent ganglion cell responses, we used the multielectrode array to stimulate several nearby ganglion cells simultaneously. From these data, we conclude that electrical stimulation of mammalian retina with small-diameter electrode arrays is achievable and can provide high temporal and spatial precision at low charge densities. We review previous epiretinal stimulation studies and discuss our results in the context of 32 other publications, comparing threshold parameters and safety limits.

Action Potentials↗

Correlated firing of cat retinal ganglion cells. I. Spontaneously active inputs to X- and Y-cells.

1. The shared inputs to cat retinal ganglion cells have been investigated by studying correlations in the maintained firing of neighboring ganglion cells. The firing of one cell was recorded from its axon in the optic tract, while that of a neighboring cell was simultaneously recorded with a second electrode in the retina. The recorded cells were of the X- or Y-type and viewed a uniform screen having a luminance of 10 cd/m2. 2. Ganglion cells with overlapping receptive-field centers showed two basic forms of correlated firing: if they had the same center sign (both on-center or both off-center), then they tended to fire at the same time, as shown by a peak in their cross-correlogram; but if they had opposite center signs (an on- and and off-center cell), they tended not to fire at the same time, as shown by a well, or dip, in their cross-correlogram. 3. Both of these tendencies were strongest for cells that were close together and did not appear for cells with nonoverlapping receptive-field centers. The strongest correlations were between neighboring Y-cells, cells with large fields, and the weakest were between X-cells, cells with small fields. In general, the strength of the correlations depended primarily on the area of the overlap between fields. 4. These correlations in maintained firing appear to be principally or entirely caused by shared inputs to the ganglion cells from more distal retinal neurons. The signals from these distal neurons appear to have strong, brief (4-8 ms), well-defined effects on ganglion cells, which are observed even in the absence of a visual stimulus. The inputs responsible for the correlated firing are thus referred to as spontaneously active inputs or simply as active inputs. 5. An analysis of the features in the various types of cross-correlograms supports the following statements about these spontaneously active inputs. a) There are two types of active inputs: inputs excitatory to on-center cells and simultaneously inhibitory to off-center center cells and inputs excitatory to off-center cells and simultaneously inhibitory to on-center cells. b) The active inputs of each type provide excitation to both X- and Y-cells of one center sign and inhibition to both X- and Y-cells of the other center sign. There is no evidence for a special class of more selective inputs providing input only to X-cells or only to Y-cells. c) Active inputs account for the majority (about 80%) of the spikes in the maintained activity of Y-cells but only a small fraction (about 15%) of the spikes in the maintained activity of X-cells. 6. A likely source of the active input signals appears to be spiking amacrine cells with a low rate of spontaneous activity.

Animals↗

Cat retinal ganglion cell receptive-field alterations after 6-hydroxydopamine induced dopaminergic amacrine cell lesions.

Optic tract single-unit recordings were used to study ganglion cell response functions of the intact cat eye after 6-hydroxydopamine (6-OHDA) lesioning of the dopaminergic amacrine cell (AC) population of the inner retina. The impairment of the dopaminergic AC was verified by high pressure-liquid chromatography (HPLC) with electrochemical detection of endogenous dopamine content and by [3H]dopamine high-affinity uptake; the dopaminergic ACs of the treated eyes demonstrated reduced endogenous dopamine content and reduced [3H]dopamine uptake compared with that of their matched controls. Normal appearing [3H]GABA and [3H]-glycine uptake in the treated retinas suggests the absence of any nonspecific action of the 6-OHDA on the neural retina. The impairment of the dopaminergic AC population was found to alter a number of response properties in off-center ganglion cells, but this impairment had only a modest effect on the on-center cells. An abnormally high proportion of the off-center ganglion cells in the 6-OHDA treated eyes possessed nonlinear, Y-type receptive fields. These cells also possessed shift-responses of greater than normal amplitude, altered intensity-response functions, reduced maintained activities, and more transient center responses. Of the on-center type cells, only the Y-type on-center cells were affected by 6-OHDA, possessing higher than normal maintained activities and altered intensity-response functions. The on-center X-cells were unaffected by 6-OHDA treatment. The dopaminergic AC of the photopically adapted cat retina therefore modulates a number of ganglion cell response properties and within the limits of this study is most prominent in off-center ganglion cell circuitry. When functioning normally, the dopaminergic AC of the cat's retina appears to make the receptive field of the off-center cell more sustained and may make its spatial summation characteristics more linear while adjusting the intensitive properties of neurons in both the on- and off-center pathways.

Animals↗

Neurotransmitter inputs to directionally sensitive turtle retinal ganglion cells.

Synaptic drugs were superfused into turtle eyecup preparation while recording extracellularly from directionally sensitive (DS) retinal ganglion cells. As in previous experiments in intact rabbit retina, both picrotoxin (a GABA antagonist) and physostigmine [an acetylcholine (ACh) potentiator] reduced or eliminated the directional selectivity of these cells. These drug effects occurred at micromolar concentrations and were long lasting. Superfusion of ACh caused excitation, and GABA caused inhibition of the spike activity of these DS cells. In some experiments, the ganglion cell was isolated from its presynaptic inputs by perfusing with a low-Ca2+/EGTA perfusate, which blocked synaptic transmission but did not suppress spike firing. During this synaptic block, ACh still caused spontaneous spike firing, and GABA was able to suppress the ACh-induced spike activity. Strychnine slightly increased the spontaneous activity of DS ganglion cells and reduced their response to light. Glycine and taurine were equally effective in totally suppressing spike activity, and strychnine blocked this inhibition by both agents. However, these inhibitory effects may be transynaptic because glycine did not suppress ACh-induced excitation during synaptic block. Superfusion of micromolar concentrations of methionine enkephalin and [D-Ala2]methionine enkephalinamide occasionally caused small increases in the light responses of DS cells, whereas naloxone, a broad-spectrum opiate antagonist, moderately decreased light responsiveness. Because naloxone had no effect on these cell's directional tuning, the opiate system is probably not involved in the mechanism of directional sensitivity. Based on the effects of these transmitter candidates and their antagonists, a possible site fo DS subunits may be the ACh and GABA receptors on the membrane of DS ganglion cells. ACh provides light-evoked excitation that may, when potentiated by physostigmine, overcome asymmetric GABA inhibition. Although the role of glycine in directional sensitivity is small, it may be responsible for regulating presynaptic excitatory pathways leading to the DS ganglion cells.

Animals↗

X-cells in the cat retina: relationships between the morphology and physiology of a class of cat retinal ganglion cells.

1. The morphology of 21 physiologically characterized X-cells in the cat retina was studied using intracellular recording and injection with horseradish peroxidase. The data from these experiments were used to test directly the relationships between specific structural and functional characteristics of a sample of individual retinal ganglion cells of the same anatomical and physiological class. Where possible, the response properties of 53 other retinal X-cells that were not successfully injected and recovered are compared with those of the labeled sample. These comparisons, which included conduction velocities (both intraretinal and extraretinal) and receptive-field size, indicate that the labeled X-cells are a representative sample of the population of retinal X-cells at corresponding eccentricities. 2. The somata of this group of injected retinal X-cells increase in size with increasing distance from the area centralis up to 13 degrees eccentricity (the greatest distance from the area centralis at which an X-cell was injected and recovered). The soma sizes of this sample of retinal ganglion cells range from 143.5 to 529.9 micron 2 (diam = 13.5-26.0 micron). Comparison of the soma sizes of the injected and recovered retinal X-cells with those of 300 Nissl-stained neurons at comparable eccentricities in the same retinae indicate that the injected sample had soma sizes that are consistent with their classification as "medium-sized" retinal ganglion cells (5, 69, 74). 3. All of the physiologically characterized retinal X-cells of this study have the compact dendritic arbors described to the morphological class of retinal ganglion cell called beta-cells by Boycott and Wässle (5). The dendrites of some of these neurons have many spinelike appendages, whereas those of other cells are nearly appendage free. We found no obvious correlation between the presence of dendritic appendages and any specific response characteristic ("ON-" or "OFF-center", etc). Like the size of the soma, both the diameter of the dendritic arbors of these cells, and the number of primary dendrites (those dendrites that originate directly from the soma), increase with increasing distance from the area centralis. 4. Since both morphological and physiological data were obtained for these neurons, it is possible to describe the relationship between the size of the receptive-field center and the diameter of the dendritic arbor for individual retinal ganglion cells. These comparisons show that the relationship between the anatomical measure and this response parameter for the entire sample of labeled X-cells is not as strong as had previously been suggested.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗