SUMMARY1. Cat retinal ganklion cells may be subdivided into sustained and transient response-types by the application of a battery of simple tests based on responses to standing contrast, fine grating patterns, size and speed of contrasting targets, and on the presence or absence of the periphery effect. The classification is equivalent to the 'X'/' Y' (linear/nonlinear) subdivision of Enroth-Cugell & Robson which is thus confirmed and extended.2. The sustained/transient classification applied to both on-centre and off-centre cells.3. Lateral geniculate neurones may be similarly classified by the same tests. Occasional concentrically organized cells had a mixture of sustained and transient properties.4. A technique for simultaneous recording from a geniculate neurone and one or more retinal ganglion cells providing its excitatory input showed that the connexions were specific with respect to the sustained/ transient classification as well as the on-centre/off-centre classification. Most geniculate neurones are excitatorily driven only by retinal ganglion cells of the same functional type. In a few cases the inputs were mixed but only with respect to the sustained/transient classification.5. Sustained retinal ganglion cells had slower-conducting axons than the transient type. The same was true for lateral geniculate neurones but in this case the distributions showed considerable overlap.6. The sustained/transient classification is the functional correlate for the well-known segregation of optic nerve fibres into two conduction groups.7. The pathways carrying sustained and transient information remain essentially separate from retina through the lateral geniculate nucleus to the striate cortex.

1. Two groups of interneurons that are involved in the organization of the lateral geniculate nucleus (LGN) are described. The cell bodies of one group lie within the LGN; these units are referred to as intrageniculate. The cell bodies of the other group are found immediately above the LGN at its border with the perigeniculate nucleus; these units are referred to as perigeniculate. 2. Intrageniculate interneurons have center-surround receptive fields that resemble those of relay (principal) cells. They can be subdivided into brisk or sluggish and sustained or transient categories. They are stimulated transsynaptically from the visual cortex and have a characteristic variation in the latency of their spike response to such stimulation both at threshold and for suprathreshold stimuli. The pathway for this stimulation appears to be via cortical efferents to the LGN. Intrageniculate interneurons receive direct, monosynaptic retinal inputs, as determined by recording simultaneously from such interneurons and from the ganglion cells which provide excitatory input to them. Similar to relay cells, they are shown to have one or two major ganglion cell inputs. 3. Perigeniculate interneurons are generally binocularly innervated and give on-off responses to small spot stimuli throughout their receptive field. They respond well to rapid movement of large targets. They respond to electrical stimulation of the retina with a spike latency that falls between that of brisk transient and brisk sustained relay cells. This latency is one synaptic delay longer than that of brisk transient relay cell activation and suggests that they are excited by axon collaterals of these relay cells. Electrical stimulation of the visual cortex is also consistent with this model; the latency of the response of perigeniculate interneurons is approximately one synaptic delay longer than the latency of the response of brisk transient relay cells. 4. The interneuronal pathways described are consistent with proposed circuits that subserve the generation of IPSPs that arise in response to optic nerve and visual cortical stimulation. We now show that such inhibition has feed-forward (intrageniculate) and feed-back (perigeniculate) components that are mediated by two different classes of geniculate interneurons. It is suggested that the intrageniculate interneurons are involved in precise, spatially organized inhibition and that the perigeniculate interneurons are part of a more general, diffuse inhibitory system that modulates LGN excitability.

The inner plexiform layer of human, monkey, cat, rat, rabbit, ground squirrel, frog and pigeon retinas was studied by electron microscopy. All showed the same qualitative synaptic arrangements: bipolar cells made dyad ribbon synapses onto amacrine and ganglion cells; amacrine cells made conventional synaptic contacts onto bipolar, ganglion and other amacrine cells. Montages of electron micrographs through the full thickness of the inner plexiform layer were made for each species and were scored for synaptic contacts. Both absolute and relative quantitative differences were found between species. The ratio of amacrine cell (conventional) synapses to bipolar cell (ribbon) synapses, the absolute number of amacrine cell synapses and the number of inter-amacrine cell synapses were all found to be higher in those animals which are known to have relatively complex retinal ganglion cell receptive field properties. It is suggested that the amacrine cell is involved in mediating complex visual transformations in certain vertebrate retinas.Physiological studies have shown that in some vertebrates the ganglion cells of the retina are relatively simple analysers of local intensity differences within the visual field. In others, many ganglion cells respond selectively to specific features of thc visual stimulus such as the size, velocity, and direction of motion of an object. Thus to varying degrees among different species, the retina acts as more than a simple transducer of patterns of light and dark.Qualitative light and electron microscopy show that all vertebrate retinas are similarly organized into three nuclear layers and two plexiform layers. The types of synaptic interconnections between cells are likewise similar. Yet it is reasonable to expect that the intricacy of the neural circuits within a retina is related to the degree of complexity of the response patterns of its ganglion cells. Recent comparative electron microscopic studies in primates and in the frog have led Dowling ('68) to suggest that these relations may be seen through quantitative studies of the synaptic patterns of the inner plexiform layer of the retina. This layer (see fig. 15) contains the axonal endings of the bipolar cells, the dendritic endings of the ganglion cells and all of the synaptic endings of the amacrine cells (Cajal, '11; Dowling and Boycott, '66; Boycott and Dowling, '69).Kidd ('62) was the first to describe inner plexiform layer synapses. He enumerated four types : conventional, spine, ribbon and serial. Conventional and spine synapses are both identified by a combination of membrane and cleft specialization, possible pre-and postsynaptic thickenings and a dense aggregation of synaptic vesicles clustered near the presynaptic membrane (see fig. 1). These are the structural attributes now widely accepted as identifying sites of neural communication in the central nervous system (Gray and Guillery, '66). In primates, Dowling and Boycott ('66) showed that both these types of synaptic contact were only made presynaptically by am...

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