Our eye has evolved to compensate for these many flaws. Our nerve cells compare the signals from the cones to measure the amount of green-or-red and blue-or-yellow, particularly along the edges of an image. In the diagram below, the signal begins with the cones, on the right. Lee BB, Smith VC, Pokorny J, Kremers J. Rod inputs to macaque ganglion cells. Vision Res. 1997;37:2813–2828. [PubMed]
. Stimuli consisting of small spots evoke spectral responses consistent with the action of only a single spectral type of cone in each case. Surround responses are also spectrally selective, and are generated only from spectral types of cone not found in the center responses (De Monasterio and Gouras, 1975). Tonic cells are spectrally opponent types, but the opponent signals are distributed in different regions of space.Evidently circuitry was the source of a spatial weighting function, shaped so that sensitivity diminished radially with distance from the ‘receptive field center’. Hartline also noted other features suggestive of hidden complexity in this simple concept. Response wave forms tended to become more transient and less sustained as spots were displaced (Fig. 5). Subtle movements, on the order of a few µm, much smaller than the receptive field itself, could evoke vigorous discharges, a kind of ganglion cell ‘hyperacuity effect’ (see also Shapley and Victor, 1986).Raynauld J-P. Goldfish retina: sign of the rod input in opponent color ganglion cells. Science. 1972;177:84–85. [PubMed]Hartline HK. The response of single optic nerve fibers of the vertebrate eye to illumination of the retina. Am J Physiol. 1938;121:400–415.
. It is a useful tool for detecting small signals. This is seen in Fig. 33 for a brief, dim stimulus which delivers on average only 5 quanta to the cornea each flash. Owing to optical losses, perhaps only 1 of these quanta is effectively absorbed in a rod and transduced into neural activity. Even so, on average, a substantial burst in mean firing rate occurs. Through analysis of data such as these Barlow et al. (1971) concluded that a single quantum absorption resulted in the firing of 2-3 extra ganglion cell nerve impulses. Cross correlations in firing patterns of adjacent cells under conditions of dim illumination agree with this estimate, and suggest the time course of increased firing probability for a single quantum absorption may last about 50 msec or more and that several adjacent ganglion cells detect the event (Mastronarde, 1983b). A retinal ganglion cell is a type of neuron located near the inner surface of the retina of the eye. For faster navigation, this Iframe is preloading the Wikiwand page for Retinal ganglion cell Mariani AP. Association amacrine cells could mediate directional selectivity in pigeon retina. Nature. 1982;298:654–655. [PubMed] While many retinal ganglion cells (RGCs) lack the spatiotemporal averaging capacity required to track changes in background light (7), ipRGCs express sustained responses to steady levels of illumination..
Retinal ganglion cells are commonly classified as On-center or Off-center depending on whether Citation: Geffen MN, de Vries SEJ, Meister M (2007) Retinal Ganglion Cells Can Rapidly Change.. Daw NW, Pearlman AL. Cat colour vision: evidence for more than one cone process. J Physiol. 1970;211:125–137. [PubMed] [Free Full text in PMC]Author contributions: N.M., R.S., and R.J.L. designed research; N.M., R.S., and C.G.E. performed research; R.S. and A.C. analyzed data; and N.M., R.S., R.S.P., and R.J.L. wrote the paper.Shapley RM, Victor JD. Nonlinear spatial summation and the contrast gain control of cat retinal ganglion cells. J Physiol. 1979;290:141–163. [PubMed] [Free Full text in PMC]
Maturana HR, Frenc S. Directional movement and horizontal edge detectors in the pigeon retina. Science. 1963;142:977–979. [PubMed] Why all the fuss? The purpose of the ganglion cells is not fully known, but they are involved in color vision. Ganglion cells compare signals from many different cones. Hartline HK. The effects of spatial summation in the retina on the excitation of the fibers of the optic nerve. Am J Physiol. 1940;130:700–711.In retinal ganglion cells, separate rod and cone pathways converge. One might expect then, during dark adaptation, as input signals to ganglion cells shift from cone-related to rod-related pathways, that receptive field properties of ganglion cells might change. The changes are, in fact, rather subtle and require careful probing with physiological stimuli and pharmacological tools. The size of receptive field center mechanisms frequently increases. Receptive field surround responses are often less vigorous. The pharmacology of light responses changes. All these effects result from changes in the dominant retinal circuitry pattern driving ganglion cells as the retina adjusts to dim lighting conditions.To evaluate coding efficiency, we estimated the information per spike in bits per spike (for T = 200 ms, ∆t = 50 ms) as Ispk = I(S; R)/(T*<firing rate>), where <.> denotes averaging across trials and time windows.
The radiance measurements shown in Fig. 1 A and B were obtained by using a custom-built portable device. Raw readings were obtained with a linear photodiode (Thorlabs). Data were sampled at 5 Hz, and acquisition was controlled by an Arduino UNO board (Arduino LLC). Radiance measurements were then calibrated by using a LIFX Color 1000 bulb.Retinal sections from hM3Dq and control OPN4Cre/+ mice were stained for c-Fos (Fig. 3D and SI Appendix, Fig. S1 A and B). There were significantly more c-Fos–positive nuclei in both the GCL and inner nuclear layer (INL) of hM3Dq-expressing retinas (Fig. 3 C and E; two-tailed unpaired t tests, P = 0.0028 and P = 0.0071 for GCL and INL, respectively). All mCherry-positive (hM3q-expressing) cells were also c-Fos–positive, but c-Fos was not restricted to these transduced ipRGCs, as for every mCherry-positive cell there were approximately eight mCherry-negative c-Fos–positive cells across INL and the GCL. These findings indicate that chemogenetically activated ipRGCs in turn excite many other cells in the GCL and INL (SI Appendix, Fig. S1C). In mice, both amacrine and ganglion cells can appear in both of these layers. To determine the degree to which the c-Fos signal appeared in ganglion cells, we counterstained with an RGC marker [RNA-binding protein with multiple splicing (RBPMS) (22)]. This revealed that the vast majority (at least 90%) of c-Fos–positive cells were indeed ganglion cells (Fig. 3F and SI Appendix, Fig. S1 A and B).In cells with input pathways arising from more than one spectral class of cone, the relative strength of different cone inputs can be modified by colored backgrounds (Wagner et al, 1960). This is called ‘selective chromatic adaptation.’ In the cell of Fig. 17, red backgrounds reduce the strength of OFF inhibitory responses, allowing green ON excitatory responses to be seen at long wavelengths. Conversely green backgrounds reduce the strength of ON responses and allow red OFF responses to be seen at short wavelengths. Modification of spectral properties by colored backgrounds is often used to evaluate cone inputs, even in the absence of color opponent signals.Central illumination of an ON-center ganglion cells depolarizes the cell membrane. Much of the normally inside negative resting potential is lost. The action excites nerve impulse activity. There is an increase in membrane conductance with a reversal potential near zero mV, typical of an open ion channel selectively permeable to cations. This synaptic mechanism causes light responses to increase in amplitude as the cell is hyperpolarized by extrinsic current. The picture is consistent with the activation of ionotropic glutamate receptors permeable to sodium, potassium, and perhaps calcium ions (Belgum et al, 1982, Freed and Nelson, 1994). In the cat ON-beta ganglion cell (Fig. 46) reversal potentials were found to shift to more negative values over the time course of the response. This appears to result from delayed activation of inhibitory synaptic currents with negative reversal potentials. Such inhibition arises in part from the receptive field surround, but also may be intrinsic to the center mechanism. A simple circuit involving excitatory ionotropic glutamate impingement from narrow field bipolar cells, and a more protracted GABAergic and/or glycinergic input from broader field amacrine cells serves to model such results. ON-center ganglion cells may receive tonic inhibitory input in the dark as well (Belgum et al, 1982).Primate retinal ganglion cells occur in two broad categories: ‘tonic’ and ‘phasic’ as described by Gouras (1968). Tonic cells respond to light stimuli in a steady maintained manner. Receptive field centers are extremely small, about 15 µm on the retinal surface (~4′ of arc, about 10 cm at 100 m distance). Tonic cells are often called ‘midgets’ because they probably represent recordings of midget ganglion cells described by Polyak (1941), see below). They occur with a density distribution across the retina comparable to anatomically midget cells (Gouras, 1968). The tiny receptive field centers match well the anatomical sizes of these tiny cells. The optic nerve conduction velocities of tonic fibers are slow (~2 m/s, Gouras, 1969). As a group these cells are also often referred to as the ‘parvocellular pathway’ or ‘P’ cells. This is because the tonic fibers terminate in the parvocellular layer of the lateral geniculate nucleus of the thalamus.
Wagner HG, MacNichol EF, Wolbarsht ML. Thee response properties of single ganglion cells in the goldfish retina. J Gen Physiol. 1960;43:45–62. [PubMed]The melanopsin ganglion cels are also thought to be involved in the pupillary light reflex for they also project to the lateral geniculate nucleus through the olivary pretectal nucleus (OPN) and on to the Edinger Westaphal nucleus (EW) for control of the pupillary light reflex (Fig. 48, light blue pathway).Only time invariant background stimulation was used in the previous two studies. When localized flashing stimuli are used, a further cross correlation pattern emerges. Oscillatory firing patterns become entrained among cells responding to a contiguous stimulus.luminance and color opponent channels, respectively. Color opponency was a color vision theory of the 19th century physiologist Ewald Herring (1875). Retina operationally combines both Young and Herring ideas in processing spectral information.
Lettvin JY, Maturana HR, McCulloch WS, Pitts WH. What the frog’s eye tells the frog’s brain. Proc Inst Radio Eng N Y. 1959;47:1940–1951.The bath applied, inhibitory retinal neurotransmitters GABA and glycine potently evoke inhibitory currents in ganglion cells. GABA receptors in cat beta cells are of the ‘A’ type. These responses can be largely blocked by the GABA antagonist bicuculline. Strychnine effectively blocks glycine-evoked currents (Cohen et al, 1994).To determine the extent to which individual units displayed a systematic change in firing rate across the ramp, we next calculated goodness of fits (R2) for a log:linear relation between firing rate and irradiance for single units. High R2 values indicate progressive changes in firing, and low values, either minimal or more discontinuous/stochastic variations in activity. Units with high R2 values were strongly biased toward increases in firing across the ramp (Fig. 2F; ON: P ∼ 0, zval = 16.25, n = 297; OFF: P ∼ 0, zval = 13.67, n = 212; sign test for change in firing in either ON or OFF units with R2 > 0.5). Indeed, of 406 ON units in these recordings, 297 had R2 > 0.5 and 289 of these increased firing; while of 303 OFF units, 212 had R2 > 0.5 and 206 of those increased firing.Barlow HB, Levick WR, Yoon M. Responses to single quanta of light in retinal ganglion cells of the cat. Vision Res. 1971;3:87–101. [PubMed]Gouras P. Antidromic response of orthodromically identified ganglion cells in monkey retina. J Physiol. 1969;204:407–419. [PubMed] [Free Full text in PMC]
Neuenschwander S, Singer W. Long-range synchronization of oscillatory light responses in the cat retina and lateral geniculate nucleus. Nature. 1996;379:728–733. [PubMed]Cross correlations are generated from simultaneous recording of 2 ganglion cell spike trains. Impulses from the first cell set the zero time around which impulse firing rate histograms for the second cell are generated. The histogram is accumulated for every impulse fired by the first cell. When two neighboring ON-center cells are so paired, a large central peak in firing rate for the second ON cell is seen (Fig. 41, red , left). A similar pattern is seen when 2 OFF cells are recorded (Fig. 41, blue right). In both cases increased firing rate represents an increased likelihood that if one cell fires, the other will also. Impulse generation in the two cells is not independent. The cells share common excitatory inputs sources, and are likely to be simultaneously excited. There is symmetry about the zero time axis. This is consistent with the idea that the cells share one or more common excitatory inputs, and that either one may respond first.
Loss of surround response can be reversed by pharmacological intervention (Fig. 35). Introduction of a cyclic AMP analogue (CPT-cAMP) into the perfusion media of this dark adapted ganglion cell restores surround responses (magenta). In further experiments (Jensen, 1991) strychnine, an antagonist of the inhibitory neurotransmitter glycine, was also found to restore surround responses to dark adapted ganglion cells. It appears thatsurround responses are actively inhibited in dark adapted retina by activation of glycinergic circuitry, probably from rod dominated amacrine cells. This glycinergic amacrine release appears modulated by the intracellular messenger cAMP.DeVries SH, Baylor DA. An alternative pathway for signal flow from rod photoreceptors to ganglion cells in mammalian retina. Proc Natl Acad Sci U S A.1995;92:10658–10662. [PubMed] [Free Full text in PMC]This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10.1073/pnas.1810701115/-/DCSupplemental.
Daw NW. Colour-coded ganglion cells in the goldfish retina: extension of their receptive fields by means of new stimuli. J Physiol. 1968;197:567–592. [PubMed] [Free Full text in PMC]Grzywacz NM, Tootle JS, Amthor FR. Is the input to a GABAergic or cholinergic synapse the sole asymmetry in rabbit’s retinal directional selectivity? Vis Neurosci.1997;14:39–54. [PubMed]This approach is similar to that described by Chichilnisky (24), the only difference being in the additional parameter “B” that we used to account for stimulus-unrelated baseline activity.Grzywacz NM, Amthor FR. Facilitation in ON-OFF directionally selective ganglion cells of the rabbit retina. J Neurophysiol. 1993;69:2188–2199. [PubMed]Shapley R, Victor J. Hyperacuity in cat retinal ganglion cells. Science. 1986;231:999–1002. [PubMed]
Belgum JH, Dvorak DR, McReynolds JS. Sustained synaptic input to ganglion cells of mudpuppy retina. J Physiol. 1982;326:91–108. [PubMed] [Free Full text in PMC]Fig. 49. Drawing and photograph of a melanopsin ganglion cell in wholemount mouse retina. The axon is indicated by an arrow. After Berson, 2003Mastronarde DN. Correlated firing of cat retinal ganglion cells. I. Spontaneously active inputs to X- and Y-cells. J Neurophysiol. 1983;49:303–324. [PubMed]Michael CR. Receptive fields of single optic nerve fibers in a mammal with an all-cone retina. II. Directionally selective units. J Neurophysiol. 1968;31:257–267.[PubMed]
Experiments were in accordance with the UK Animals (Scientific Procedures) Act 1986 approved by Home Office (PPL Number 70/8918) and performed on adult (3–6 mo) Opn4Cre/+;Z/EGFP mice housed in a 12-h dark/light cycle at 22 °C with food and water available ad libitum.Westhiemer G. The spatial sense of the eye. Invest Ophthalmol Vis Sci. 1979;18:893–912. [PubMed]
Hartline introduced the ‘nearly definable’ concept of ‘receptive field’ to describe the spatial properties of retinal ganglion cells. He used ‘spot mapping’ to define such fields, a technique still widely employed. Cells were found to respond to relatively dim spots when the stimulus was positioned in the ‘center’ of the receptive field but brighter stimuli were required as the spots were moved away from this region. An example of the ‘spot mapping’ technique, taken from Kuffler (1953) shows the reduction in response vigor as stimulus spots are displaced from the center (Fig. 5). Hartline concluded that ganglion cell receptive fields were fixed in space and immobile, typically did not extend beyond 1 mm in diameter, and were graded in sensitivity over this region. Receptive fields were much larger than expected of individual photoreceptors, suggesting signal processing and integration through retinal circuitry.Reid RC, Shapley RM. Spatial structure of cone inputs to receptive fields in primate lateral geniculate nucleus. Nature. 1992;356:716–718. [PubMed]Wyatt HJ, Daw NW. Specific effects of neurotransmitter antagonists on ganglion cells in rabbit retina. Science. 1976;191:204–205. [PubMed]Barlow HB, Hill RM, Levick WR. Retinal ganglion cells responding selectively to direction and speed of image motion in the rabbit. J Physiol. 1964;173:377–407.[PubMed]
Abbreviations: RGC, retinal ganglion cells; Shh, sonic Hedgehog; FGF8, fibroblast growth factor 8; DAPT, (N-[N-(3,5-difluorophenacetyl)-l-alanyl]-S-phenylglycine t-butyl ester); BDNF.. Review and cite RETINAL GANGLION CELLS protocol, troubleshooting and other methodology information | Contact experts in RETINAL GANGLION CELLS to get answers
Ganglion cell axons terminate in brain visual centers, principally the lateral geniculate nucleus and the superior colliculus. Ganglion cell axons are directed to specific visual centers depending of the visual ‘trigger features’ they encode. The optic nerve collects all the axons of the ganglion cells. In man this optic nerve bundle contains more than a million axons.Daw NW, Pearlman AL. Cat colour vision: One cone process or several? J Physiol. 1969;201:745–764. [PubMed] [Free Full text in PMC]We find that gradual increments in ambient light (irradiance ramps) are indeed accompanied by increases in RGC firing and that this is associated with an increase in information rate. The change in firing occurs in absence of any other visual stimulus and is partially disrupted in the absence of melanopsin (the photopigment of ipRGCs), indicating that it is a genuine change in intrinsic activity as a response to alterations in ambient light, and at least partly attributable to ipRGCs. Chemogenetic activation of ipRGCs recapitulates increases in both firing and information rates. This latter finding not only confirms the importance of ipRGCs in setting RGC firing, but establishes a causative relationship between firing and information rates by showing that increasing firing is sufficient to enhance information in the absence of any change in the visual environment. These data reveal a mechanism for increasing visual information at higher ambient light and establish the potential for proactive control of neuronal firing to be used to scale information flow in the nervous system according to predictable changes in demand.There are 8 synaptic ion channel associated retinal neurotransmitter receptors . Such receptors are referred to as ‘ionotropic’. Glutamate, the most common excitatory neurotransmitter, activates three distinct channels: AMPA type channels, kainate type channels, and NMDA type channels. Acetyl choline is a further excitatory neurotransmitter active at nicotinic receptors. Excitatory receptors are associated with channels selective for positively charged ionic species, monovalent or divalent cations. When open they allow positive charges to enter neurons, depolarizing them and exciting impulse firing. GABA and glycine are inhibitory ionotropic neurotransmitters. GABA activates two distinct ionotropic receptor types: the bicuculline sensitive GABAA receptor and the bicuculline insensitive GABAC receptor. Bicuculline is a convulsant, and a common blocker of GABA inhibitory action. GABA and glycine open channels selective for the negatively charged chloride anion, which hyperpolarizes neurons and inhibits impulse firing. In teleost fish retinas a further inhibitory chloride selective channel present in bipolar cells has been found activated by glutamate. All these ionotropic receptors are heteropolymers composed typically of 5 subunits. There are multiple alleles of these subunits allowing for considerable heterogeneity in receptor composition and details of function.APB may not effectively block rod signals in all ganglion cell types however. DeVries and Baylor (1995) showed In rabbit retina, that rod responses in ‘brisk’ ganglion cells were eliminated by APB, but rod responses of ‘directionally selective’ or ‘sluggish’ ganglion cell types were not. Rod signals that could not be blocked in these ganglion cells may use a pathway through rod-cone photoreceptor gap junctions into the APB-resistant OFF cone bipolar pathway.
As noted by Hartline, not only strength of response, but also a change in form of response occurs as stimuli are moved across ganglion cell receptive fields. Kuffler (1953) noted a particularly abrupt change in form, from onset excitation for stimulation in the center of the field to offset excitation for stimulation in the receptive field periphery. An example is illustrated in Fig. 6. In Fig. 6a, a spot, centered near the electrode tip, evokes a burst of impulses at stimulus onset. In Fig. 6b the spot is displaced by 0.5 mm, where it evokes no impulses at onset, but a burst of impulses at offset. In Fig. 6c an intermediate spot position is found where both actions are evoked by the stimulus. The example (Fig. 6) shows properties of a classic ON-center, receptive field, with OFF surround. Conversely OFF-center cells, which are excited at stimulus offset by central stimulation, have regions in peripheral receptive field where ON excitation is evoked. This is the classic center-surround receptive field organization of ganglion cells. Such organization was further examined with intracellular recording techniques. These techniques revealed not only the ganglion cell impulses, but also the slow changes in ganglion cell membrane potential which modulate the impulse rate.Given the finite duration of our WN sequence, the space of all possible stimuli cannot be explored during the experiment. Therefore, it is possible that the particular pseudorandom sequence of our illumination values provides a bias itself in the estimate of the amount of information conveyed by individual units. To control for this possibility in CNO experiments, we divided each trial in two separated epochs of temporal white noise. We then separately calculated mutual information for each epoch. The results for each unit were well matched (SI Appendix, Fig. S2E), indicating that the duration of each stimulation epoch is enough to capture its coding properties.Ganglion cells are the final output neurons of the vertebrate retina. Ganglion cells collect information about the visual world from bipolar cells and amacrine cells (retinal interneurons). This information is in the form of chemical messages sensed by receptors on the ganglion cell membrane. Transmembrane receptors, in turn, transform the chemical messages into intracellular electrical signals. These are integrated within ganglion-cell dendrites and cell body, and ‘digitized’, probably in the initial segment of the ganglion-cell axon, into nerve spikes. Nerve spikes are a time-coded digital form of electrical signalling used to transmit nervous system information over long distances, in this case through the optic nerve and into brain visual centers.To further evaluate the ability of individual neurons to represent an incoming stimulus, in addition to the previously described measure of mutual information, we used a decoding approach. We divided the stimulation sequence into two epochs that we used respectively for estimating the LNP model (training set) and perform maximum a posteriori decoding (test set). Decoding performances on the test set were then evaluated by measuring Pearson’s linear correlation between the original and the reconstructed stimulus (ρorig-rec in Figs. 5 G and H and 6 G and H).
Noise in the visual signal falls as ambient light increases, allowing the retina to extract more information from the scene. We show here that a measure of ambient light produced by the small number of inner retinal photoreceptors [intrinsically photosensitive retinal ganglion cells (ipRGCs)] regulates intrinsic rates of spike firing across the population of retinal ganglion cells that form the optic nerve. Increased firing at higher irradiance allows the ganglion cells to convey more information. Our findings reveal a potential mechanism for increasing visual performance at high ambient light and show that changes in maintained activity can be used to provide proactive control over rates of information flow in the CNS.CLICK HERE to see an animation of the center and surround receptive field organization of a beta ganglion cell (Quicktime movie)
Retinal Pigment Epithelium, Retina (Ophthalmology). Peripapillary Retinal Nerve Fiber Layer and Ganglion Cell-Inner Plexiform Layer Thickness in Children with Familial Mediterranean Fever where r represents the neural response and N represents the number of time windows in each trial. The response r was calculated in a time window of duration T as the L-dimensional vector of firing rates r = [r[0, ∆t), r[∆t,2*∆t), …, r[(l − 1)*∆t, L*∆t)] within L adjacent subwindows of duration ∆t such that T = L*∆t. The response entropy is determined by the time-averaged distribution of firing rates P(r) = <P(r|t)>t, where P(r|t) is the probability of response r in the time window [t − T/2, t + T/2) and <.>t denotes averaging across time windows. For a neuron that is insensitive to changes in illumination, then P(r|t) = P(r) at any given time t and the mutual information will be zero.
While much of the analysis presented here concerns the consequences of changes in maintained activity for the transfer of higher-frequency visual signals, it is certainly possible that irradiance-driven changes in firing rate are themselves also a method of conveying information. There is a growing body of evidence that neurons can convey multiple types of information by multiplexing across different timescales (6, 35). In this context, the same RGCs could convey information about visual patterns in the fine timing of spikes, and irradiance in their maintained activity. The latter irradiance code appears in the thalamus (9) and could be exploited at higher levels to support perception of ambient light (36) or as contextual information for visual prediction and for associative learning (37).Amthor FR, Takahashi ES, Oyster CW. Morphologies of rabbit retinal ganglion cells with complex receptive fields. J Comp Neurol. 1989;280:97–121. [PubMed]Cleland BG, Dubin MW, Levick WR. Sustained and transient neurones in the cat’s retina and lateral geniculate nucleus. J Physiol. 1971;217:473–496. [PubMed] [Free Full text in PMC]
Cronly-Dillon JR. Units sensitive to direction of movement in goldfish optic tectum. Nature. 1964;203:214–215. [PubMed]Phasic cells combine red and green cone signals synergistically in both center and opponent surround mechanisms of the receptive field. These cells are luminosity types and are not generally color opponent, although in some cases an imbalance of the strength of spectral mechanisms in center and surround generates some opponent characteristics (De Monasterio and Gouras, 1975).STAs were calculated for each isolated unit. For the ramp protocol, we calculated six STAs per unit, each corresponding to a half log unit increment in irradiance. For the WN+CNO experiments, the STAs were separately calculated before and after CNO delivery. Each STA was tested (t test; α = 0.01) for significance by comparing its energy with a null distribution obtained by randomly shuffling the times of spike occurrence along the time of the recording (20 repeats). All significant STAs (n = 6,983) were standardized and projected along the first five principal components (the number of components was determined as the smallest set that could explain >75% of the STA variance). We then applied a kmeans clustering for two groups to separate units into ON and OFF polarity (Fig. 2D).Central illumination of OFF-center ganglion cells hyperpolarizes the cell membrane. The normally inside negative cell potential increases in magnitude. This action reduces nerve impulse activity and causes the cell to become quiet. Concomitantly there is an increase in membrane conductance with a reversal potential more negative than the resting potential. This is typical of the opening of channels selectively permeable to the chloride anion. (Belgum et al, 1982). A simple circuit involving narrow-field GABA or glycine releasing amacrine cells which are themselves excited by light serves to model the OFF center response.
The neurotransmitter acetyl choline is a potent excitatory agent for directionally selective ganglion cells (Ariel and Daw, 1982). It is released by presynaptic cholinergic ‘starburst’ amacrine cells and provides excitation at both onset and offset of the stimulus. Blockade of cholinergic input (Cohen and Miller, 1995; Grzywacz et al, 1997), or ablation of cholinergic amacrine cells, reduces but does not abolish directional selectivity (He and Masland, 1997). The metabotropic glutamate receptor agonist APB blocks the ON component of all responses in ON-OFF type directionally selective cells (Cohen and Miller, 1995; Kittila and Massey, 1995). However, the OFF component of directional responses remains. NMDA (N-methyl-D-aspartate) receptor agonists do not block directionally selective responses, and by implication directionally selective ganglion cells contain excitatory NMDA response components which are non-directional (Cohen and Miller, 1995). Interestingly, the AMPA-kainate receptor antagonist NBQX potently blocks directional selectivity in both ON-OFF and ON type cells leaving vigorous responses to movement in all directions (Fig. 16). Thus a non-NMDA glutamate receptor may be selectively involved in the directionally selective mechanism, possibly as an excitatory (bipolar) input to a GABAergic interneuron (Cohen and Miller, 1995).The average firing rate of each unit was estimated in of the 20 WN epochs before and after CNO delivery. We then tested each unit for the possibility that CNO induces a systematic change in average firing by performing an unpaired rank sum between firing-rate distributions measured before and after CNO delivery. Category:Retinal ganglion cells. From Wikimedia Commons, the free media repository. Jump to navigation Jump to search Rod system delays occur in specialized retinal circuitry devoted to rods. The retina has developed a set of neurons and synapses especially tuned for processing visual signals under dim lighting conditions. These may include specialized synapses between rods and bipolar cells, as well as specialized sets of rod amacrine cells. These cells are tuned to the longer integration times of rods.Yang G, Masland RH. Direct visualization of the dendritic and receptive fields of directionally selective retinal ganglion cells. Science. 1992;258:1949–1952.[PubMed]
X and Y cells are differentiated in other ways as well. X cells are numerous and have rather narrow receptive fields, and in cat retina are utilized for highest acuity vision. These are morphologically the ‘beta’ cells of cat retina. Y-cells are rather sparsely distributed and have wide receptive fields. These are morphologically the ‘alpha’ cells of the cat, and most likely serve a visual alerting function. They are sometimes named according to the shape of the response wave form to steps of light. Y cells are called ‘brisk transient’ cells, and X cells are called ‘brisk sustained’ cells (Cleland and Levick, 1974a). Y cells are also distinguished from X cells by larger axon diameters and quicker conduction times from retina to the thalamic visual centers of the brain. These sets of cells initiate two separate and parallel pathways for transfer of visual information to brain visual centers. (Cleland et al, 1971). Morphological features and the distribution of X/beta, Y/alpha ganglion cells are discussed below.As a first test of the hypothesis that ipRGCs contribute to irradiance-dependent changes in GCL activity, we asked whether this behavior was disrupted in melanopsin-knockout mice (Opn4−/−). Because ipRGCs integrate information from rods and cones with their own melanopsin-dependent intrinsic light response, irradiance responses are typically disrupted rather than abolished following melanopsin loss (9, 14, 15). This turned out also to be the case for the change in GCL firing. As firing activity in dimmest condition was different between these strains (0.35 ± 0.04 SEM, 0.49 ± 0.05 SEM; P ∼ 0, zval = 6.442; n = 662 and 455, rank sum test; Opn4−/− and control animals), we compared the change in firing rate across the ramp. While the irradiance ramp did elicit an increase in firing in Opn4−/− retinas, the magnitude of this effect was significantly reduced compared with that of melanopsin-sufficient controls (Fig. 3A).Enroth-Cugell C, Robson JG. The contrast sensitivity of retinal ganglion cells of the cat. J Physiol. 1966;187:517–552. [PubMed] [Free Full text in PMC]
Fig 15A illustrates the static receptive field properties of an ON directionally selective cell in rabbit retina. This cell responds to spots moving towards the lower left (P), but not the upper right (N, Fig 15B). No responses are seen to bars of light, regardless of the direction of motion. This indicates a particularly strong surround antagonism suppressing responses to large stimuli. Picrotoxin abolishes both directional selectivity and size specificity. In these experiments (Wyatt and Daw, 1976) strychnine, another convulsant and blocker of the inhibitory neurotransmitter glycine,did not have these effects. This suggested that GABA was selectively important in neural circuitry underlying the generation of directionally selective responses. The GABAA selective antagonist SR95531 blocks directional responses suggesting that this particular subtype of GABA receptor is critical to the process (Kittila and Masey, 1995).Your input will affect cover photo selection, along with input from other users. Listen to this article Thanks for reporting this video!Retinal ganglion cells respond to all common excitatory or inhibitory retinal neurotransmitters. When neurotransmitters are applied to the solution bathing ganglion cells, membrane currents are induced. AMPA, kainate (Fig. 44) or NMDA evoke excitatory currents in both ON and OFF type cat beta cells (Cohen et al, 1994). NMDA currents are of the typical ‘conditional’ sort, dominant only if cells are depolarized first by other excitatory neurotransmitters, or in the absence of extracellular magnesium (Fig. 45). Extracellular acetylcholine also excites retinal ganglion cells (Masland and Ames, 1976; Lipton et al, 1987; Cohen et al, 1994).
Retinal Ganglion Cells play a key role in the visual system, being the only connection between the retina and the areas of the brain dedicated to the process of the visual information Jensen RJ. Involvement of glycinergic neurons in the diminished surround activity of ganglion cells in the dark-adapted rabbit retina. Vis Neurosci. 1991;6:43–53.[PubMed]Impact of ipRGC-driven increases in firing on neural coding. (A) As in Fig. 5A for a representative CNO-responsive unit from an hM3Dq-expressing retina; arrow to Right shows time of CNO administration. (B and C) Trial-to-trial reproducibility (B) and information rate (C) for the block of 40 trials after (“CNO”) than before (“pre”) CNO delivery in units expressing increase in firing rates post-CNO (P = 0.00014, P ∼ 0 for G and B, P = 0.433 and 0.229 for S and T; n = 393; sign test). (D–H) As in Fig. 5 D–H, but applied to CNO+WN experiments in hM3Dq retinas; accordingly, distribution of Δρorig-red in H is for neurons exhibiting CNO-related increases in firing. (I) Coding efficiency, calculated as bits per spike, was equivalent in “dim” vs. “bright” portions of the irradiance ramp (Left) but significantly reduced following CNO delivery (Right). ***P < 0.001, ****P < 0.0001 for rank sum test. ganglion cell - a nerve cell whose body is outside the central nervous system; damage to Klein, Gene-regulation logic in retinal ganglion cell development: Isl1 defines a critical branch distinct from..
Unlike the phasic cell (Fig. 26) this blue-excited cell is able to respond to color changes, being excited by a change from yellow to blue stimuli of equal luminance (Fig. 28c). As a bistratified type it is clearly not a midget ganglion cell. Dacey and Lee plausibly argue that the cell receives OFF type red and green bipolar cell input onto its outer, OFF-layer dendritic tree, and ON-type blue bipolar cell input onto its inner, ON-layer dendritic tree. Dacey and Lee also describe a small, peripheral monostratified ganglion cell responding to red-green color changes, but not to the magenta-green color changes designed to excite blue sensitive ganglion cells. As yet foveal and parafoveal midget cells have not been marked by microelectrode staining techniques.Fig. 11b. The null test for linear summation within X- and Y-type receptive fields (Enroth-Cugell and Robson, 1966) Retinal Ganglion Cells: Anatomy. Ganglion cell dendrites extend into the inner plexiform layer (IPL), a neuropil located on the outer side of the ganglion cell layer Symmetry about the time zero axis suggests that in fact two symmetric inputs are required: One inhibiting the ON cell while exciting the OFF cell, and the other exciting the ON cell while inhibiting the OFF cell. Physiologically ON and OFF ganglion cell mosaics are not independent then. Mutual inputs are found. Such inputs occur, however, only for cells with overlapping, or partially overlapping receptive fields, as is also the case for paired ON or paired OFF cells (Mastronarde, 1983a).Edited by Fred Rieke, University of Washington, and accepted by Editorial Board Member Jeremy Nathans November 1, 2018 (received for review June 21, 2018)
Purpura K, Kaplan E, Shapley RM. Background light and the contrast gain of primate P and M retinal ganglion cells. Proc Natl Acad Sci U S A. 1988;85:4534–4537. [PubMed] [Free Full text in PMC]Brivanlou IH, Warland DK, Meister M. Mechanisms of concerted firing among retinal ganglion cells. Neuron. 1998;20:527–539. [PubMed]
But receptive field size is controlled by more than the retinal ganglion cell (in particular, horizontal and amacrine cells) which have wider and wider umbrellas to collect signals in the periphery The retinal ganglion cells (RGCs) are the output cells of the retina into the brain. In mammals, these cells are not able to regenerate their axons after optic nerve injury, leaving the patients with optic.. Barlow HB, Levick WR. The mechanism of directionally selective units in the rabbit’s retina. J Physiol. 1965;178:477–504. [PubMed]
Chemogenetic activation of ipRGCs enhances maintained firing across the RGC population. (A) Schematic of our visual-chemogenetic stimulus. A white-noise (WN) stimulus was presented at constant mean irradiance (11.5 log10 photons⋅cm−2⋅s−1) with timed CNO delivery. (B) Representative image for a whole-mount retina rec orded under our MEA system. mCherry fluorescence spanning large regions of the recording array could be observed. (C) Mean ± SEM change in time-averaged firing rate (ΔFR over mean before CNO administration) for hM3Dq-expressing (blue line) and control (black line) retinas following CNO application (5 µM) commencing at time 0. The top blue-black dashed line indicates significant differences in ΔFR (**P < 0.01, rank sum tests, Bonferroni’s correction). (D) The distribution of logarithmic ratios between firing rates (log10frratio) from hM3Dq-expressing retinas, before and after CNO delivery, is unimodal, and most units express an increase in firing rate (log10frratio > 0). (E) As in B, plotting ON and OFF units from hM3Dq retinas (lines above indicate significant increases in firing; P < 0.01, sign rank tests, Bonferroni’s correction). (F) Pie charts presenting percentage of units that showed increase, decrease, or no change in firing rate upon CNO application in hM3Dq (Left) and control (Right) retinas.Tonic cells of the parvocellular pathway respond best to stimuli with high contrast and fine grain, while phasic cells to stimuli with very weak contrast covering larger areas (Kaplan and Shapley, 1986).Color opponent ganglion cells can have centers and surrounds each with separate color opponent properties. A common type in goldfish retina is the double opponent ganglion cell. In this type each color mechanism in the receptive field center is opposed by a mechanism of the same color type but opposite sense in the surround. Spectral properties of a goldfish ganglion cell with a red OFF center, red ON surround in combination with a green ON center, green OFF surround appears in Fig. 20. This type of cell is thought to excel at ‘simultaneous color contrast’: the ability to detect color changes at borders.Müller F, Wässle H, Voigt T. Pharmacological modulation of the rod pathway in the cat retina. J Neurophysiol. 1988;59:1657–1671. [PubMed]Torstein Wiesel, 1981 winner of the Nobel prize in physiology and medicine, provides an example of such an intracellular recording in Fig. 7, which allows further features of the surround response to be seen. In Fig. 7a, a small spot of light depolarizes the membrane of the cell and evokes a vigorous burst of impulses. This is an ON-center cell. In Fig. 7b the stimulus is an annulus, a ring of light designed to stimulate the receptive field surround, but not the center. Impulses are silenced during stimulus presentation, but a burst appears at annulus offset. This is a classic surround pattern. A hyperpolarization of the membrane occurs while the annulus is ON and impulses are silenced. This is a feature of the response hidden from extracellular recording methods. The surround stimulus actively inhibits the cell by hyperpolarizing its membrane. Finally, the response to the large spot stimulus (Fig. 7c) should not be neglected. The large spot, like the small spot (Fig. 7a) evokes a membrane depolarization and sustained burst of impulses. But the depolarization is smaller, and the burst less vigorous. The net impact of the center-surround receptive-field structure is that ganglion cells ‘prefer’ small spots to large spots! Size selectivity is a unique and telltale characteristic of ganglion cell physiology. To summarize, ganglion cell center-surround interaction is a multifaceted phenomenon:
Some pharmacological agents abolish, or partially abolish directional selectivity. The most classic of these is picrotoxin, a convulsant, and blocker of receptors for the inhibitory neurotransmitter gamma-aminobutyric acid (GABA). Both GABAAand GABAC type receptors, the latter very common in retina, are blocked by picrotoxin.Ganglion cells respond to colored stimuli in one of two ways: color opponent responses and luminance responses. The patterns are similar to those of horizontal cells, which occur in chromatic and luminance types. Color opponent ganglion cells are found principally in vertebrates which behaviorally can discriminate color. Goldfish are an animal model with color vision and have provided much information on the way in which vertebrate retinas process color. In this animal it was first realized that individual cones express only one each of three genomically available cone photopigment types (colloquially red, green, or blue cones, technically long, L, mid, M, or short, S, wavelength types). This supported at least part of Thomas Young’s 19th century ‘trichromatic theory of color vision. Neural retina does not preserve the purity of thisspectral trilogy, however. Signals are both added and subtracted to formCohen ED, Zhou ZJ, Fain GL. Ligand-gated currents of alpha and beta ganglion cells in the cat retinal slice. J Neurophysiol. 1994;72:1260–1269. [PubMed] Other articles where Ganglion cell is discussed: human eye: The retina: innermost layer of neurons, the ganglion cells; and the transmitted messages are carried out of the eye along their projections.. Ex vivo retinas were illuminated using a combination of violet, blue, cyan, and yellow (λmax at 400, 430, 480, and 560 nm, respectively) elements of a multispectral LED light source (SPECTRA X light engine; Lumencor). The intensity distribution of the temporal white-noise modulations were as described previously (9) and produced by pulse width modulation of LEDs controlled via the counter channels of a NI USB-6343 card and LabVIEW software (National Instruments Corporation). The irradiance ramp was produced by a gradient neutral-density filter wheel (Newport Corporation) controlled by a stepper motor (Newport Corporation) to provide 5.12 log-equally spaced steps per second, each step corresponding to 0.2% increase in irradiance. The filtered light was then collected by a liquid light guide and delivered to the mouse retina mounted on the MEA. The effective irradiance for each photopigment was calculated as previously (33).
To determine what fraction of units displayed a significant increase in firing, we compared mean firing rates over 40 repeats of the white-noise stimulus on either side of CNO administration for each single unit. Thirty-one percent of units from hM3Dq retinas (122/393 units) showed a statistically significant increase in firing post-CNO, with only 1.5% (6/393 units) showing a significant reduction over this time frame (Fig. 4F; rank sum test with Bonferroni-corrected α = 0.05). In the control retinas, the figures were 7.7% and 3% (24/309 and 9/309 units, respectively; Fig. 4F). Furthermore, a comparison of firing-rate profiles for the 20% units with highest increase in activity following CNO in the two groups revealed a substantially larger increase in the hM3Dq population (SI Appendix, Fig. S2A). Thus, not only were increases in firing more common in hM3Dq retinas, but, when present, their amplitude was qualitatively different.Schmidt M, Humphrey MF, Wässle H. Action and localization of acetylcholine in the cat’s retina. J Neurophysiol. 1987;58:997–1015. [PubMed]Information rate is defined as limT→∞I(S;R)/T. In practice, we estimated the information rate, in bits per second, as the slope of I(S;R) as function of T (T = 50, 100, 150, and 200 ms), given a fixed temporal subwindow (∆t = 50 ms).
Retinal ganglion cells in the largest biology dictionary online. Free learning resources for students covering all major areas of biology In Fig. 32 turtle ganglion cell impulses are evoked by electric current injection directly into rods and cones themselves. The delay to onset of the first impulse discharge in underlying ganglion cells is measured. This technique separates circuitry delays from delays in photoreceptor transductive machinery. OFF discharge latency (the time between cessation of hyperpolarizing current injection into a photoreceptor and the first ganglion cell impulse) was found to be about 100 msec when electrically stimulating red cones, but a longer (150 msec) when stimulating rods. Interestingly ON-discharges evoked by red cone stimulation (time from hyperpolarizing current onset to first spike) were also slow (170 msec), suggesting a slower pathway for ON cone information as well, possibly through a metabotropic glutamate pathway in such cone bipolar cells.Daw NW, Ariel M. Effect of synaptic transmitter drugs on receptive fields of rabbit ganglion cells. Vision Res. 1981;21:1643–1648. [PubMed]Receptive fields and dendritic fields are similarly spaced. Multielectrode arrays allow simultaneous recording of all ganglion cell light responses in a patch of retina, characterization of trigger features, and mapping of receptive fields (DeVries and Baylor, 1997). Such studies show that receptive fields of ganglion-cell types having common trigger features approach each other only at the edges. When spatial profiles are fit with 2 dimensional Gaussian functions, receptive field centers are typically 2 Gaussian radii apart. In this arrangement, summation of sensitivities from all cells of a given type leads to a flat sensory surface (Devries and Baylor, 1997).
Fig. 7. Changes in ganglion cell membrane potential accompanying center and surround responses (Wiesel, 1959,)Nelson R, Kolb H, Freed M. Off-alpha and Off-beta ganglion cells in cat retina. I. Intracellular electrophysiology and HRP stains. J Comp Neurol. 1993;329:68–84.[PubMed] Ganglion cells are the final output neurons of the vertebrate retina. Ganglion cells collect information about the visual world from bipolar cells and amacrine cells (retinal interneurons) Hartline’s electrical recordings of single optic nerve fiber responses revealed ‘discharges of impulses’, or ‘action potentials’ in response to light stimulation. The discharge patterns were, however, diverse (Hartline, 1938). Three unique patterns of light response were described (Fig.3).In addition to information about color, ganglion cells also transmit to the brain a monochromatic, non color containing signal from rods. Rods detect visual stimuli at very low light levels, including visual threshold. Their true sensitivity is rarely tested in a typical urban environment, but imagine a moonless rural night without street lights. Shapes and forms are fuzzy, colorless, and slowly perceived. This is rod vision. Such visual environments are often termed ‘scotopic’, in contrast to brighter ‘photopic’ environments where shape color and movement are readily seen. In photopic environments cones are active. Amazingly, individual ganglion cells transmit both sorts of information, a sort of multiplexing of signals with very different characteristics, though it is only with a ‘mesopic’ visual environment, with intermediate levels of ambient illumination, that both signals may be simultaneously present.
Dear Viewers of these Videos- These lectures are from my undergrad course The Human Brain, currently being taught in the spring of 2018 at MIT Rod signals develop more slowly than cone signals. Reaction times driving automobiles or playing sports out of doors is slower at night. This is because rod signals are characteristically slow and it is this photoreceptor system that is active at night or with dim illumination.Given a constant and relatively large window T, both P(r) and P(r|t) are estimated on subwindows of finite duration ∆t and the choice of ∆t will affect the estimate of mutual information. Although ideally we would like ∆t to be as small as possible, this would result in a very high-dimensional response vectors that cannot be properly sampled with a finite number of stimulus trials (see Bias correction). On the other hand, an extremely conservative choice of ∆t will fail to capture the temporal precision of the neural responses. To identify a suitable trade-off, we first choose a value of T = 200 ms because comparable with the latency of the STAs peaks (Fig. 1G). We then calculated the mutual information by changing the number of subwindows L. As mutual information tended to saturate with L (SI Appendix, Fig. S2D), we selected ∆t = 50 ms (corresponding to L = 4), which provided a good estimate of the temporal precision for neural responses in our data.In Fig. 29, extracellular spikes from a primate ON-center retinal ganglion cell can be seen composed of two distinct clusters with different delays. With yellow backgrounds only short latency spikes are generated in response to a brief flash, regardless of wavelength or stimulus brightness. These are cone signals. Under dark adapted conditions and threshold stimulation, only long latency spikes occur. These are rod signals. Under dark adaptation and with somewhat brighter red stimuli, which begin to stimulate cones, a mixed response can be seen composed of both short latency cone components (arrow) and long latency rod components. In Fig. 30 rod and cone responses are seen in intracellular recordings of a cat OFF-center ganglion cell. Responses to flickering red (647 nm) stimuli arise mainly from red cones, while those from blue (441 nm) stimuli arise only from rods. Superposition of responses allows easy comparison of the timing of rod and cone signals with each flicker cycle. Rod ON hyperpolarizations evoked by the blue stimulus are distinctly delayed as compared to cone ON hyperpolarizations evoked by the red stimulus. The different delay is also apparent with the excitatory impulses evoked at the offset phase of each cycle.
Intracellular recordings from phasic ganglion cells in isolated monkey retina (Fig. 26) confirm that these responses originate in ‘parasol’ type ganglion cells (Fig. 27). The phasic, parasol response is chromatically a luminosity type. Vigorous responses are evoked by alternating bright and dark periods (Fig. 26a) but not by alternating blue and yellow colors (Fig. 26b), or magenta and green colors (Fig. 26c) balanced for brightness. Phasic cells sense luminance changes but not color changes (Dacey and Lee, 1994).Mice were held in the dark for 12 h (to enhance integrity of dissected retina) before cervical dislocation under dim red light (50 nW⋅cm−2, λ > 650 nm). Still working under dim red light, retinas were isolated in carboxygenated (95% CO2, 5% CO2) aCSF, with concentrations (in mM): 118 NaCl, 25 NaHCO3, 1 NaH2PO4, 3 KCl, 1 MgCl2, 2 CaCl2, 10 C6H12O6, and 0.5 l-glutamine (Sigma-Aldrich). The retina was incised radially multiple times along the edge to maximize planarization and then mounted onto a 256-channel MEA (256MEA200/30iR-ITO; Multichannel Systems) with the GCL facing down onto the electrodes. A Cyclopore membrane (5-μm pores; Whatman) held the retina in place while being weighed down by two stainless-steel anchors (∼0.75 g each) bearing a framework of parallel polyimide-coated fused silica capillaries (TSP320450; Polymicro Technologies).Barlow HB, Hill RM. Selective sensitivity to direction of movement in ganglion cells of the rabbit retina. Science. 1963;139:412–414. [PubMed]