In cat retina, rod bipolar terminials do not synapse on ganglion cells but on two types of amacrine cell (types I and II). Cone bipolars synapse directly on ganglion cells and on type I amacrines. The type II amacrine appears to play a special internuncial role between bipolars and ganglion cells in the rod system.
The inner plexiform layer of the mammalian retina has a bisublaminar organization determined by restricted branching of the terminals of cone bipolar cells and dendrites of class I (large) and class II (small) ganglion cells. Comparison of dendritic field diameters and receptive fiedl center sizes of large ganglion cells suggests that neural circuitry in sublamina a conveys "OFF"-center properties and connections in sublamina b "ON"-center properties to retinal ganglion cells.
1. Ganglion cells in the retina of the cat were stained by intracellular dye injection after recording their responses to photic stimulation. 2. All cells encountered were divided into those giving on-responses and those producing off-responses, and the level of dendritic branching of these two groups was compared. Cells giving off-responses were found to branch high in the inner plexiform layer (IPL), near the amacrine cell bodies (sublamina a); those giving on-responses were found to branch lower in the inner plexiform layer (sublamina b). 3. Dye-injected cells varied widely in morphology and size, having cell bodies ranging in diameter from 8 to 32 micrometer and dendritic fields ranging from 25 to 490 micrometer in diameter; yet the sign of the response of each unit correlated only with the level of dendritic branching. Thus, no other morphological feature except stratification appears to be important in determining the sign of the response of these cells. 4. The stratification of ganglion cells into on- and off-layers parallels the distribution of the axon terminals of the flat and invaginating cone bipolars. Flat cone bipolars are in a position to contact off-center ganglion cells (in sublamina a) and invaginating cone bipolars are in a position to contact on-center ganglion cells (in sublamina b). 5. The rod and cone inputs to some cells were characterized by comparing their responses to deep red and blue rod-matched stimuli over a 2-log unit range starting at dark-adapted threshold. About half the cells appeared to be rod dominated under these conditions, whereas the others appeared to have mixed rod and cone signals. 6. The nature of the rod and cone pathways to ganglion cells is discussed.
The synaptic organization of starburst amacrine cells was studied by electron microscopy of individual or overlapping pairs of Golgi-impregnated cells. Both type a and type b cells were analyzed, the former with normally placed somata and dendritic branching in sublamina a, and the latter with somata displaced to the ganglion cell layer and branching in sublamina b. Starburst amacrine cells were thin-sectioned horizontally, tangential to the retinal surface, and electron micrographs of each section in a series were taken en montage. Cell bodies and dendritic trees were reconstructed graphically from sets of photographic montages representing the serial sections. Synaptic inputs from cone bipolar cells and amacrine cells are distributed sparsely and irregularly all along the dendritic tree. Sites of termination include the synaptic boutons of starburst amacrine cells, which lie at the perimeter of the dendritic tree in the "distal dendritic zone." In central retina, bipolar cell input is associated with very small dendritic spines near the cell body in the "proximal dendritic zone." The proximal dendrites of type a and type b cells generally lie in planes or "strata" of the inner plexiform layer (IPL), near the margins of the IPL. The boutons and varicosities of starburst amacrine cells, distributed int he distal dendritic zone, lie in the "starburst substrata," which occupy a narrow middle region in each of the two sublaminae, a and b, in rabbit retina. As a consequence of differences in stratification, proximal and distal dendritic zones are potentially subject to different types of input. Type b starburst amacrines do not receive inputs from rod bipolar terminals, which lie mainly in the inner marginal zone of the IPL (stratum 5), but type a cells receive some input from the lobular presynaptic appendages of rod amacrine cells in sublamina a, at the border of strata 1 and 2. There is good correspondence between boutons or varicosities and synaptic outputs of starburst amacrine cells, but not all boutons gave ultrastructural evidence of presynaptic junctions. The boutons and varicosities may be both pre- and postsynaptic. They are postsynaptic to cone bipolar cell and amacrine cell terminals, and presynaptic primarily to ganglion cell dendrites. In two pairs of type b starburst amacrine cells with overlapping dendritic fields, close apposition of synaptic boutons was observed, raising the possibility of synaptic contact between them. The density of the Golgi-impregnation and other technical factors prevented definite resolution of this question. No unimpregnated profiles, obviously amacrine in origin, were found postsynaptic to the impregnated starburst boutons.(ABSTRACT TRUNCATED AT 400 WORDS)
Bipolar, amacrine, and ganglion cells of carp retina, stained intracellularly with Procion yellow, can be divided into types a and b, according to the destination of terminals and dendritic trees in the inner plexiform layer (sublamina a and b, respectively). Type a cells showed hyperpolarizing, or off, responses and type b cells depolarizing, or on, responses. Carp thus resembles cat in the basic organization of on and off pathways in the retina.
The morphological characteristics of Class 111, Golgi Type I1 neurons are analyzed in Golgi impregnations of the lateral geniculate nucleus (LGN) in the cat with special attention given to the complex dendritic appendages of these cells, the multilobed dendritic terminals. In addition, electron microscopic studies demonstrate the presence of presynaptic dendritic (ID) terminals as major components of synaptic glomeruli. Electron microscopic analysis of serial thin sections confirm the dendritic origin and the complex, multilobed nature of these terminals. An hypothesis is proposed identifying the ID terminal with the multilobed dendritic terminals of light microscopically identified Class I11 cells and the IA terminal, the "peripheral axon" of the glomerulus, with the axonal endings of all Golgi Type I1 cells.The Class I11 neuron has a short axon ramifying near the cell of origin. The axon terminates in boutons en passant and boutons terminaux: single, bilobed or "claw-like" endings. Optic axons terminate not in "claw-like" fashion, as previously thought, but in club-shaped endings with scalloped silhouettes. The principal dendrites of Class I11 cells are characterized by long-stalked appendages, often multilobed and occurring at the end of preterminal stalks 5 to 100 p in length. One impregnated multilobed terminal studied in the electron microscope was found to innervate two adjacent glomeruli.The relationship of ID terminals to optic axon terminals and to other processes in synaptic glomeruli is analysed with the aid of the electron microscope. The ID terminal contains vesicles of the same size as those in presynaptic dendrites and makes small, macular synaptic contacts with other pre-and postsynaptic dendrites. ID terminals are the only elements in the LGN which synapse on others of their own kind, and are postsynaptic to all types of axon identified. As the only postsynaptic processes containing synaptic vesicles in the dorsal layers of the LGN, they support the conclusion that there are no axo-axonic synapses in these layers. The ID terminal, the optic terminal and spinous dendrites originating presumably from geniculo-cortical relay (GCR) cells, are seen to enter into a triadic relationship characteristic of the synaptic glomerulus in which the presynaptic dendrite synapses on the spinous dendrites and the optic terminal synapses on both dendrites. ID terminals reconstructed from serial sections communicate as "internuncial" presynaptic dendrites between optic central axons in adjacent synaptic glomeruli. IA terminals, which like ID terminals contain flattened vesicles, are presynaptic to ID terminals, as well as to postsynaptic dendrites, cell somata and axon initial segments.A model for the internuncial character of multilobed dendritic terminals and the potential for inhibition of Golgi Type I1 axon terminals is proposed, and theories about the function of Gold Type I1 cells in lateral geniculate transmission are briefly considered.The synaptic glomerdus The electron microscopic appearance OP th...
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