Interplexiform Cells Research Articles

Glycinergic synaptic inputs to bipolar cells in the salamander retina.

1. Glycine activated strychnine-sensitive chloride conductances at both the dendrites and the axonal telodendria of most bipolar cells in the salamander retina. 2. The chloride equilibrium potential of bipolar cells was found to be negative to -50 mV, indicating that glycinergic synapses on bipolar cells are inhibitory. 3. Some bipolar cells exhibited discrete, strychnine-sensitive, chloride-mediated inhibitory postsynaptic currents (IPSCs). These were elicited by focal application of glutamate at the inner plexiform layer (IPL). Glycinergic synapses were localized using simultaneous focal application of calcium to retinal slices bathed in calcium-free media. Both dendritic and telodendritic glycinergic IPSCs were observed. 4. The decay of the telodendritic IPSCs was well fitted by a single exponential with a time constant of 17.7 +/- 8.7 ms. Similar kinetics were observed for dendritic IPSCs in some cells, but in one class of on-centre bipolar cell the decay of the dendritic IPSCs was better fitted by a sum of two exponentials with time constants 9.9 +/- 4.3 and 51.3 +/- 24.3 ms. 5. The dendritic IPSCs were best driven by application of glutamate at the distal IPL (the off sublamina), while the telodendritic IPSCs were driven best by application near the telodendria. These results suggest that bipolar cell dendrites receive inhibitory glycinergic inputs from interplexiform cells that are excited by off-centre bipolar cells, whereas bipolar cell telodendria receive glycinergic amacrine cell inputs that are antagonistic to the photoreceptor inputs. 6. Both inputs could be elicited in the presence of tetrodotoxin (TTX), but the dendritic IPSCs were sometimes abolished by TTX, suggesting that sodium-dependent spikes play an important role in the transmission of interplexiform cell signals to the outer plexiform layer.

Theory and implementation of infomax filters for the retina

In the first part of this paper we discuss a technical visual sensory system, which—in analogy with the retina—includes some preprocessing of visual information. In so doing, we use an information-theoretic criterion, the infomax ansatz, to optimize the response of the sensory system. In particular, it is shown that the lattice structure of the photoreceptor array has to be taken into account. By a discrete Fourier transformation on a triangular lattice we derive the frequency response of the infomax filter within the first Brillouin zone. To illustrate the response properties, infomax filters adapted to different noise levels are applied to images with different signal-to-noise ratios. This clearly demonstrates the necessity of adaptation of the filter properties to the given noise level. Furthermore, it is shown how to efficiently implement infomax-like filters by simple networks with only nearest-neighbour interactions. A two-layered network topology proves to be very advantageous in implementing the desired high-pass or low-pass properties. The network topology allows for adaption of the network to low and high noise levels by simply adjusting the nearest-neighbour couplings.In the second part of this paper, we compare the previously described information-theoretic requirements on a visual sensory system with biological facts known from the vertebrate retina. The substantial physiological response properties of the vertebrate retina are in agreement with the main features of the infomax filter. Since available experimental data lacks information which is necessary for a more quantitative comparison, we present suggestions for future experiments. Some key anatomical features of the retina of many vertebrates compare well with our two-layered implementation of the infomax filter. The analogy, in particular, concerns the adaption mechanism. To illustrate this point, we summarize some recent experiments which demonstrate that in the retina of some species adaption is based on the release of the neuromodulator dopamine by the interplexiform cells. This causes the horizontal cells to decouple. On the basis of recently gained understanding of the outer plexiform layer of the retina some further hypotheses about the functionality of the retina become obvious and possible future experiments to verify or refute them are suggested.Finally, we discuss the infomax approach from a more general point of view. In particular, we show that redundancy is essential to obtaining noise robustness of an internal representation of the environment as it is produced by a sensory system such as the retina.

Immunocytochemical localization of gamma-aminobutyric acid plasma membrane transporters in the tiger salamander retina.

Gamma-aminobutyric acid (GABA) plasma membrane transporters (GATs) play an important role in regulating GABA neurotransmission in the nervous system. The distribution of two GATs, GAT 1 and GAT 3, in salamander retina was investigated by using affinity-purified polyclonal antisera directed to the predicted C-terminals of rat GAT 1 and rat GAT 3. GAT 1-immunoreactivity (-IR) was found in type IB and IIB orthotopic bipolar cells (BCs) located in the distal and middle of the inner nuclear layer (INL), respectively; in type IIA and IA amacrine cells (ACs) located in the middle and proximal INL, respectively; and in interplexiform cells and cells in the ganglion cell layer (GCL). No detectable staining was found in horizontal cells (HCs) or in structures resembling Müller cells. GAT 1-immunoreactive fibers were present in the outer plexiform layer (OPL) and inner plexiform layer (IPL) in three bands corresponding to the three bands previously reported to be GABA-IR. GAT 3 antibodies labeled fewer cells and cell types than GAT 1 antibodies. GAT 3-IR was localized to type IIA and IA ACs and cells in the GCL, but not to BCs, HCs, or Müller cell-like structures. There was weak labeling of the OPL and stronger labeling of the IPL, with three distinct bands at the same depth as observed with GAT 1-IR. Double-labeling showed that the majority of GAT 1-IR BCs (88%), ACs (88%), and cells in the GCL (78%) colocalized with GABA-IR. The present study provides the first direct evidence of the expression of two GAT subtypes in neurons of nonmammalian retinas. These transporters could regulate GABA neurotransmission by reuptake and termination of GABA's action and, perhaps, by GABA release mechanisms. The presence of GAT 1-IR/GABA-IR bipolar cells further supports our earlier observations that a subgroup of orthotopic bipolar cells are likely to be GABAergic.

Tyrosine hydroxylase expression in the Cebus monkey retina.

Tyrosine hydroxylase (TH) expression was used as a marker to study the dopaminergic cells in the Cebus monkey retina. Two types of dopaminergic cells were identified by cell body size and location, level of arborization in the inner plexiform layer, and amount of immunolabeling. Type 1 cells displayed intense immunoreactivity and larger somata (12-24 microns) located in the inner nuclear layer or ganglion cell layer, whereas type 2 had smaller cell bodies (8-14 microns) found either in the inner plexiform layer or ganglion cell layer and were more faintly labeled. Interplexiform cells were characterized as type 1 dopaminergic cells. Immunoreactive axon-like processes were seen in the nerve fiber layer, and a net of fibers was visible in the foveal pit and in the extreme periphery of the retina. The population of TH+ cells was most numerous in the temporal superior quadrant and its density peaked at 1-2 mm from the fovea. Type 1 TH+ cells were more numerous than type 2 cells at any eccentricity. Along the horizontal meridian, type 1 cell density was slightly higher in temporal (29 cells/mm2) than in nasal (25 cells/mm2) retina, while type 2 cells had a homogeneous distribution (4.5 cells/mm2). Along the vertical meridian, type 1 cells reached lower peak density (average 17.7 cells/mm2) in the inferior retina (central 4 mm), compared to the superior portion (23.7 cells/mm2). Type 2 cell density varied from 4.5 cells/mm2 in the superior region to 9.4 cells/mm2 in the inferior region. The spatial density of the two cell types varied approximately inversely while the total density of TH+ cells was virtually constant across the retina. No correlation between dopaminergic cells and rod distribution was found. However, we suggest that dopaminergic cells could have a role in mesopic and/or photopic vision in this species, since TH+ fibers are present in cone-dominated regions like the foveola and extreme nasal periphery.

Differential reinnervation of retinal bipolar cell dendrites and axon terminals by dopamine interplexiform cells following dopamine depletion with 6-OHDA

Depletion of retinal dopamine in goldfish increases light sensitivity at photopic backgrounds. As horizontal cells appear not to be involved with this effect (Yazulla and Studholme [1995] Vis. Neurosci. 12:827-837), we investigated the innervation patterns of the ON rod/cone bipolar cells (ON-BC) by dopaminergic interplexiform cells (DA-IPCs) normally and during the period of neogeneration of new DA-IPCs at the marginal zone following DA-IPC destruction. DA-IPCs were destroyed via intraocular injection of 6-hydroxydopamine over 2 successive days. Controls and 1 year post-injection retinas were double labeled for protein kinase C and tyrosine hydroxylase (TH) immunocytochemistry to identify the ON-BCs and the DA-IPCs, respectively. Double-labeled 25 microns tissue sections were examined on a confocal laser scanning microscope by using dual channel immunofluorescence acquisition. Image stacks were analyzed for DA-IPC/ON-BC contacts in the distal inner nuclear layer (INL) and inner plexiform layer (IPL). Image stacks were rotated 180 degrees with respect to each other and reanalyzed to determine potential randomness of the contacts. For control retinas there were 1.8 contacts/axon terminal in the IPL (n = 165) and 9.4 contacts/ON-BC in the distal INL (n = 28). At 1 year after injection, reinnervation of TH-immunoreactive boutons in the retina recovered to 16% of control in the IPL but only 10% in the distal INL. Establishment of DA-IPC/ON-BC contacts recovered to 36% of control for ON-BC axon terminals (n = 103), whereas there was no recovery of contacts in the distal INL (n = 30). Reinnervation of ON-BC by DA-IPCs preferentially targets the axon terminals. The absence of reinnervation of bipolar cell dendrites by DA-IPCs may account for the persistence of the increased light sensitivity following retinal dopamine depletion. Thus, dopamine input to ON-BCs in the outer retina maybe involved in setting background sensitivity under photopic conditions.

Differential reinnervation of retinal bipolar cell dendrites and axon terminals by dopamine interplexiform cells following dopamine depletion with 6‐OHDA

Differential reinnervation of retinal bipolar cell dendrites and axon terminals by dopamine interplexiform cells following dopamine depletion with 6‐OHDA

Immunocytochemical localization of dopamine D1 receptors in the retina of mammals.

Dopamine is one of the major neurotransmitters in the retina. It is released from amacrine and interplexiform cells into both inner (IPL) and outer (OPL) plexiform layers. Several dopaminergic actions are known to occur through D1 receptors (D1R) but the precise location of these receptors has not been established. An antibody that recognizes the intracytoplasmic C-terminal of the rat D1R was used to detect D1R, immunohistochemically, in rats (Wistar and RCS), mouse, hamster, and macaque monkey retinas. The OPL was heavily stained in each species, consistent with the known actions of dopamine on horizontal cells. Three to five bands were observed in the IPL, depending on species. Three were in the a sublayer, the outermost of which was close to the amacrine cell layer, and may represent the massive dopamine input to the AII rod-amacrine cells. As observed in mice, where bipolar cells are D1-immunoreactive, the band located in sublayer 3 of the IPL may contain cone-bipolar cell terminals. A band of D1R-immunoreactivity in the b sublayer of the IPL contains ON-bipolar cell terminals and a second site of interaction between dopaminergic cells and the AII amacrine cells. This sublayer was absent from the RCS rat retina, suggesting a severe impairment of the rod-driven pathway following rod degeneration in these mutant rats. Cells in the ganglion cell layer exhibited relatively heavy staining, and may be ganglion cells or displaced amacrine cells. Some extrasynaptic localizations of D1R in the retina are suggested.

A neural model of foveal light adaptation and afterimage formation.

Psychophysical research has documented the existence of three processes in light adaptation: a fast subtractive process, a divisive process that is fast at light onset and slower at light offset, and a very slow subtractive process (Hayhoe et al., 1987). In the neural model developed here, the fast subtractive process is identified with horizontal cell feedback onto cones and the divisive process with amacrine cell feedback onto bipolar cells. The very slow subtractive process is identified with the modulatory feedback circuit from amacrines via interplexiform cells to horizontal cells. A nonlinear dynamical model is developed incorporating these aspects of retinal circuitry along with both ON- and OFF-center M and P pathways. This model is shown to account for many aspects of foveal light adaptation, including negative afterimage formation, and to explain a number of the physiological differences between M and P ganglion cells, including their differing contrast-response functions.

Dopaminergic and GABAergic retinal cell populations in mammals

A number of modern techniques now allow histologists to characterize subpopulations of retinal neurons by their neurotransmitters. The morphologies and connections of these chemically defined neurons can be analyzed precisely at both light and electron microscope levels and lead to a better understanding of retinal circuitry. The dopaminergic neurons form a loose population of special wide-field amacrine cells bearing intraretinal axons within the inner plexiform layer. One subtype, the interplexiform cell, sends an axon to the outer plexiform and outer nuclear layers. The number of interplexiform cells is variable throughout mammalian species. The GABAergic neurons form a dense and heterogeneous population of amacrine cells branching at all levels of the inner plexiform layer. The presence of GABA in horizontal cells seems to be species-dependent. Close relationships occur between dopaminergic and GABAergic cells. GABA antagonizes a number of dopaminergic actions by inhibiting both the release and synthesis of dopamine. This inhibition can be supported by GABA synapses onto dopaminergic cells, but GABA can also diffuse to its targets. Finally, GABA is also contained and synthesized in dopaminergic cells. This colocalization might be the basis of an intracellular modulation of dopamine by GABA. Microsc Res Tech 36:26–42, 1997. © 1997 Wiley-Liss, Inc.

Dopaminergic and GABAergic retinal cell populations in mammals.

A number of modern techniques now allow histologists to characterize subpopulations of retinal neurons by their neurotransmitters. The morphologies and connections of these chemically defined neurons can be analyzed precisely at both light and electron microscope levels and lead to a better understanding of retinal circuitry. The dopaminergic neurons form a loose population of special wide-field amacrine cells bearing intraretinal axons within the inner plexiform layer. One subtype, the interplexiform cell, sends an axon to the outer plexiform and outer nuclear layers. The number of interplexiform cells is variable throughout mammalian species. The GABAergic neurons form a dense and heterogeneous population of amacrine cells branching at all levels of the inner plexiform layer. The presence of GABA in horizontal cells seems to be species-dependent. Close relationships occur between dopaminergic and GABAergic cells. GABA antagonizes a number of dopaminergic actions by inhibiting both the release and synthesis of dopamine. This inhibition can be supported by GABA synapses onto dopaminergic cells, but GABA can also diffuse to its targets. Finally, GABA is also contained and synthesized in dopaminergic cells. This colocalization might be the basis of an intracellular modulation of dopamine by GABA.

Horizontal cell spinule dynamics in fish are affected by rearing in monochromatic light

Blue acaras ( Aequidens pulcher, Cichlidae) were reared for 1 yr in white or monochromatic “red”, “green” and ”bluerd lights to study the function and control mechanisms of horizontal cell (HC) spinules in the synaptic pedicles of cones. Ratios of spinules per synaptic ribbon (S/R) were determined in tangential sections in both single and double cones. We found that the S/R ratios in light adapted retinae decreased with decreasing wavelength of the rearing light in all cone types. Conversely, there was an increasing number of incompletely formed spinules with the highest frequency in the blue light group. Dark adaptation resulted in the complete degradation of mature spinules. However, significant numbers of incompletely degraded spinules were observed in the group reared in blue light. Fish reared in blue light which were transferred to white light formed mature spinules when light adapted and still had vestigial spinules when dark adapted. The mechanisms of spinule formation and degradation and the control of spinule dynamics appear to be fully developed in fish reared in monochromatic light. However, long-term chromatic deprivation seems to induce a compensatory modulation of spinule dynamics. A working hypothesis is formulated that interprets the observed effects as manifestations of differences in the activition of dopaminergic interplexiform cells (light adapted) and the sensitivity to glutamate of HCs (dark adapted). Our findings are consistent with the hypothesis that spinules are involved in signin-verting feedback transmission from HCs to cones.

A MODEL OF AN ADAPTIVE RECEPTOR BASED ON THE RETINA

A model of the retina based on biological data is developed to study the processing of images by the retina as a model for an adaptive receptor. The direct visual pathway (photoreceptors, bipolar and ganglion cells) and the horizontal units and their gap-junctions, as well as the dopaminergic interplexiform cells and dopamine release are modeled. A small network of 12 units is first simulated in order to study the behavior of the different components involved, and the role played by lateral inhibition is analyzed. A 64×64 photoreceptors model, containing 16,448 units, is then simulated. The following properties were found for stimuli that remain constant in time: (i) The difference between the time constants of the photoreceptors and the horizontal units lead to both the formation of a transient output image that contains pertinent features of the input, and the formation of a reverse after-image when the stimulus is removed. (ii) Global modifications of lateral inhibition strength allow either contrast rendering or contour enhancement. Local modifications of lateral inhibition strength bring about adaptation to local input characteristics.

Doparninergic control of light-adaptive synaptic plasticity and role in goldfish visual behavior

Dopamine has been implicated in processes of retinal light and dark adaptation. In goldfish retina, horizontal cell dendrites elaborate neurite processes (spinules) into cone terminals, in a light- and dopamine-dependent manner. However, the functions of retinal dopamine and the horizontal cell spinules in visual behavior are unknown. These issues were addressed in behavioral, electroretinographic, and anatomical studies of normal fish and those with unilateral depletion of retinal dopamine induced by intraocular (i.o.) injections with 6-hydroxydopamine (6-OHDA). Dopamine interplexiform cells (DA-IPC) disappear within 2 weeks after 6-OHDA injection; cell bodies appear at the marginal zone within 6 weeks at which time neurites slowly reinnervate the retina with a sparse plexus over the next 12 months. We found that dopamine depletion increased light sensitivity at photopic but not scotopic backgrounds by 2.5 log units, an effect mimicked by i.o. injections of dopamine DI and D2 antagonists. The ERG b-wave increment thresholds were the same for control and dopamine depleted eyes, indicating a normal transition from rod to cone systems in the ON pathway. Light-dependent spinule formation was reduced by about 60% in dopamine-depleted retinas, but returned to normal by 3 months and 9 months after injection in the entire retina, even areas not directly innervated with DA-PC processes. Spinule formation in vivo was inhibited 50% with i.o. injection of SCH 23390 in control retinas as well as throughout 3 month 6-OHDA injected retinas, including DA-IPC free areas. This latter result indicates a volume effect of dopamine, diffusing laterally through the retina over several millimeters, in regulating spinules. We conclude that DA-IPCs regulate sensitivity to background at photopic levels not via the ON pathway, but perhaps the OFF pathway. Goldfish display both increased sensitivity to light and a normal Purkinje shift in the ERG b-wave whether or not horizontal cell spinules are present, indicating that dopamine control of photopic vision in fish is not mediated through light-induced spinule formation of horizontal cell dendrites.

Multiple γ‐aminobutyric acid plasma membrane transporters GAT‐1, GAT‐2, GAT‐3 in the rat retina

γ-Aminobutyric acid (GABA) plasma membrane transporters (GATs) influence synaptic neurotransmission by high-affinity uptake and release of GABA. The distribution and cellular localization of GAT-1, GAT-2, and GAT-3 in the rat retina have been evaluated by using affinity-purified polyclonal antibodies directed to the C terminus of each of these GAT subtypes. Small GAT-1-immunoreactive cell bodies were located in the proximal inner nuclear layer (INL) and ganglion cell layer (GCL), and processes were distributed to all laminae of the interplexiform layer (IPL). Varicose processes were in the optic fiber layer (OFL) and the outer plexiform layer (OPL). Weak GAT-1 immunostaining surrounded cells in the INL and GCL, and it was found in the OFL and OPL and in numerous processes in the outer nuclear layer (ONL) that ended at the outer limiting membrane. GAT-1 is therefore strongly expressed by amacrine, displaced amacrine, and interplexiform cells and weakly expressed by Müller cells. GAT-2 immunostaining was observed in the retina pigment epithelium and the nonpigmented ciliary epithelium. GAT-3 immunoreactivity was distributed to the OFL, to all laminae of the IPL, GCL and INL, and to processes in the ONL that ended at the outer limiting membrane. Small GAT-3-immunoreactive cell bodies were in the proximal INL and GCL. GAT-3 is therefore strongly expressed by Müller cells, and by some amacrine and displaced amacrine cells. Together, these observations demonstrate a heterologous distribution of GATs in the retina. These transporters are likely to take up GABA from, and perhaps release GABA into, the synaptic cleft and extracellular space. This suggests that GATs regulate GABA levels in these areas and thus influence synaptic neurotransmission. © 1996 Wiley-Liss, Inc.

Multiple gamma-Aminobutyric acid plasma membrane transporters (GAT-1, GAT-2, GAT-3) in the rat retina.

gamma-Aminobutyric acid (GABA) plasma membrane transporters (GATs) influence synaptic neurotransmission by high-affinity uptake and release of GABA. The distribution and cellular localization of GAT-1, GAT-2, and GAT-3 in the rat retina have been evaluated by using affinity-purified polyclonal antibodies directed to the C terminus of each of these GAT subtypes. Small GAT-1-immunoreactive cell bodies were located in the proximal inner nuclear layer (INL) and ganglion cell layer (GCL), and processes were distributed to all laminae of the interplexiform layer (IPL). Varicose processes were in the optic fiber layer (OFL) and the outer plexiform layer (OPL). Weak GAT-1 immunostaining surrounded cells in the INL and GCL, and it was found in the OFL and OPL and in numerous processes in the outer nuclear layer (ONL) that ended at the outer limiting membrane. GAT-1 is therefore strongly expressed by amacrine, displaced amacrine, and interplexiform cells and weakly expressed by Müller cells. GAT-2 immunostaining was observed in the retina pigment epithelium and the nonpigmented ciliary epithelium. GAT-3 immunoreactivity was distributed to the OFL, to all laminae of the IPL, GCL and INL, and to processes in the ONL that ended at the outer limiting membrane. Small GAT-3-immunoreactive cell bodies were in the proximal INL and GCL. GAT-3 is therefore strongly expressed by Müller cells, and by some amacrine and displaced amacrine cells. Together, these observations demonstrate a heterologous distribution of GATs in the retina. These transporters are likely to take up GABA from, and perhaps release GABA into, the synaptic cleft and extracellular space. This suggests that GATs regulate GABA levels in these areas and thus influence synaptic neurotransmission.

Dopamine inhibits mammalian photoreceptor Na+,K+-ATPase activity via a selective effect on the alpha3 isozyme.

The rat retina contains dopaminergic interplexiform cells that send processes to the outer plexiform layer where dopamine is released in a light-dependent manner. We report herein that physiologically relevant concentrations of dopamine inhibited ouabain-sensitive photoreceptor oxygen consumption in dark- and light-adapted rat retinas and inhibited Na+,K+-ATPase specific activity (EC 3.6.1.37) in a rat rod outer-inner segment preparation. Experiments with the selective D1 agonist fenoldopam or D2 agonist quinpirole and experiments with dopamine plus either the D1 antagonist SCH23390 or D2/D4 antagonist clozapine showed that the inhibition of oxygen consumption and enzyme activity were mediated by D2/D4-like receptors. The amphetamine-induced release of dopamine, monitored by the inhibition of oxygen consumption, was blocked by L-2-amino-4-phosphonobutyric acid and kynurenic acid. Pharmacological and biochemical experiments determined that the IC50 values of ouabain for the alpha1-low and alpha3-high ouabain affinity isozymes of photoreceptor Na+,K+-ATPase were approximately 10(-5) and approximately 10(-7) M, respectively, and that the D2/D4-like mediated inhibition of Na+,K+-ATPase was exclusively selective for the alpha3 isozyme. The dopamine-mediated inhibition of alpha3 first occurred at 5 nM, was maximal at 100 microM (-47%), had an IC50 value of 382 +/- 23 nM, and exhibited negative cooperativity (Hill coefficient, 0.27). Prior homogenization of the rod outer-inner segment completely prevented the long-lasting inhibition, suggesting that the effect was coupled to a second messenger. Although the physiological significance of our findings to photoreceptor function is unknown, we hypothesize that these results may have relevance for the temporal tuning properties of rods.

IMMUNOHISTOCHEMICAL LOCALIZATION OF GABA IN CHAMELEON RETINA ( CHAMALEO CHAMALEO)

We used a policlonal antiserum against GABA and demonstated GABA-immunoreactivity (GABA-IR) in several populations of amacrine cells in the inner nuclear layer (INL), and other cells in the inner plexiform layer (IPL) of the central and peripheral retina of the chameleon. Horizontal cells do not contain GABA-IR and the chameleon retina is therefore an exception among non-mammals. GABA-IR was not seen in cell bodies in the position of photoreceptor, bipolar and interplexiform cells suggesting that GABA is not involved in synaptic transmission in the outer plexiform layer of chameleon retina.

Catecholaminergic interplexiform cells in the retina of lizards

An immunohistochemical study of the distribution of tyrosine hydroxylase has been performed in the retina of lizards of the genera Podarcis, Anolis and Tarentola. Immunoreactive cells extending their processes into the inner plexiform layer were observed in all three species. Reactive fibres in the outer plexiform layer were also seen in Podarcis and Anolis, and hence they possess not only amacrine but also catecholaminergic interplexiform cells. The retina of Anolis also showed reactive fibres aposed to the photoreceptors near the central fovea. The role of this outer retinal innervation on dopamine-dependent light-adaptive phenomena is discussed from a comparative perspective.

Electron microscopic demonstration of tyrosine hydroxylase-immunoreactive interplexiform cells in the lamprey retina

To clarify the controversies about the existence (or not) of dopaminergic interplexiform cells in the lamprey retina, we have performed an immunocytochemical electron microscopic study of the retina of the river lamprey Lampetra fluviatilis, using anti-tyrosine hydroxylase antibody. We demonstrate the presence of immunoreactive processes in both inner (IPL) and outer (OPL) plexiform layers. The external processes are in close contact with horizontal cell processes and photoreceptor terminals in the OPL, but do not make classical synapses. Some of them are ensheathed within Müller cell cytoplasm. Thus, dopaminergic interplexiform cells do exist in Lampetra fluviatilis, but the absence of synapses rather supports a paracrine action of dopamine in the outer retina.

Retinal neurons of the California ground squirrel, Spermophilus beecheyi: a Golgi study.

Although the optic nerve fibers of the cone-dominant ground squirrel retina have been well studied physiologically, the morphological details of the retinal neurons have not. To that end, retinal neurons of the California ground squirrel have been studied in Golgi-impregnated wholemounts. Two types of horizontal cell have been identified: H1 has an axon and axon terminal, whereas H2 is axonless. The dendritic field of H1 cells enlarges in a nonuniform manner with increasing displacement from the central retina. The smallest examples lie centrally in the visual streak, and the largest occur in the superior periphery. Eight types of bipolar cell are distinguished by morphological differences in dendritic branching pattern and field size in the outer plexiform layer, cell body size, and layering within the inner nuclear layer and by the morphology and stratification of axon terminals in the inner plexiform layer. A large bistratified bipolar cell (B8) is introduced here; the other 7 types closely resemble those in the retinas of other sciurid species described by R.W. West (1976, J. Comp. Neurol. 168:355-378; 1978, Vision Res. 18:129-136). The B1 type is proposed as a blue cone bipolar cell. Amacrine cells are classified into 27 cell types. Six of these occur as mirror-image pairs across the inner plexiform layer, the soma of one of each pair being "displaced" to the ganglion cell layer. The best described of these pairs is the very elaborate starburst amacrine cell, A5, which stains regularly in these wholemounted retinas. Changes in dendritic field size of both A5 subtypes with retinal location are quantified. The morphology of three amacrine cell types identified in Spermophilus beecheyi suggests that their possible counterparts in S. mexicanus (West, 1976) were, as displaced amacrine cells, misidentified as ganglion cells. Amacrine cell types that may play roles in the rod pathway, the blue cone pathway, and ganglion cell directional selectivity are discussed. No type of interplexiform cell was observed. Ganglion cells are classified into 19 cell types, 9 of which probably correspond to the ganglion cells described by West (1976) in the Mexican ground squirrel. The bistratified G11 cell is proposed as an ON-OFF directionally selective type.