Neurons and fibres in the chick and homing pigeon hippocampus were described following Golgi impregnation. Two principal classes of neurons were distinguished: projection neurons with distant projecting axons and spiny dendrites, and local circuit neurons. In the homing pigeon and chicken hippocampus there are three types of projection neurons: pyramidal, pyramidal-like and multipolar. The pyramidal and pyramidal-like neurons are only found in the central 'pyramidal' layer of the hippocampus whereas multipolar neurons are present in the suprapyramidal, pyramidal and infrapyramidal layers. The axon of projection neurons typically emits several varicose collaterals from the initial section. Most of these collaterals extend along the infrapyramidal layer of the hippocampus, while others ascend to the pyramidal and suprapyramidal layers where they branch. The number of impregnated axon collaterals was higher in the homing pigeon than in the chick hippocampus. A variety of multi-angular/ovoid local circuit neurons ranging from small to large size are found in the homing pigeon and chick hippocampus. Their axons develop local arborization of varicose branches, the extent of which varies with the type of local circuit neurons. The density of GABA immunopositive local circuit neurons was found to be greater in the homing pigeon than in the chick. The profuse arborization of projection neuron axon collaterals and the higher density of GABA-immunopositive local circuit neurons in the homing pigeon hippocampus may underlie the differences in hippocampal function between the homing pigeon and chick, and this complex local connectivity may contribute to the ability of spatial orientation and memory.
The neuronal structures and their possible connections in the optic tectum of chicks were studied by using Golgi impregnation, in light microscopy, and Golgi-GABA immunogold staining in electron microscopy. The terminals of the optic fibers displayed different patterns and orientation in various laminae of the avian optic tectum. The shape of the branching and the terminals of the optic fibers conformed to the postsynaptic neuronal structures. In Golgi preparations, the terminals in tectal layers 4 and 5 seemed to be located rather densely, and they contact, among other things, the ‘dendritic terminal sections’ of large ganglion cells of layer 13 of the tectum. This connection could be responsible for direct optic transmission via ganglion cells to the nucleus rotundus. Additionally, the contacts established by optic terminals in these layers with the tectal neurons arranged with their dendrites parallel to the surface of the tectum, could provide the structural basis for an inhibitory system. The terminals of isthmic nuclei were found in this optic lamina, too. The optic terminals in layer 7 contacted the dendritic side branches of some radial tectal neurons.
Following a demonstration of Golgi-impregnated neurons and their terminal axon arborization in the optic tectum, the neurons of the nucleus parvocellularis and magnocellularis isthmi were studied by means of postembedded electron-microscopical (EM) gamma-aminobutyric acid (GABA)-immunogold staining. In the parvocellular nucleus, none of the neuronal cell bodies or dendrites displayed GABA-like immunoreactivity in EM preparations stained by postembedded GABA-immunogold. However, numerous GABA-like immunoreactive and also unlabeled terminals established synapses with GABA-negative neurons. GABA-like immunoreactive terminals were usually found at the dendritic origin. Around the dendritic profiles, isolated synapses of both GABA-like immunoreactive and immunonegative terminals established glomerulus-like structures enclosed by glial processes. All giant and large neurons of the magnocellular nucleus of the isthmi displayed GABA-like immunoreactivity. Their cell surface was completely covered by GABA-like immunoreactive and unlabeled terminals that established synapses with the neurons. These neurons are thought to send axon collaterals to the parvocellular nucleus; their axons enter the tectum opticum. The morphological characteristics of neurons of both isthmic nuclei are like those of interneurons, because of their numerous axosomatic synapses with both asymmetrical and symmetrical features. These neurons are not located among their target neurons and exert their modulatory effect on optic transmission in the optic tectum at a distance.
Three types of local circuit neurons have recently been reported in the homing pigeon hippocampus. The principal type appears to be constituted by the medium-sized angular or ovoid local circuit neurons that occur in all layers of the hippocampus. The current Golgi study has revealed that these neurons can be classified according to their axonal arborisation extension: (1) in all directions, (2) principally medio-laterally, or (3) dorso-ventrally. The local circuit neurons with dorso-ventral axon arborisation are present only in the subpyramidal layer. Serial sections of a Golgi-impregnated medium-sized, multiangular local circuit neuron in the pyramidal layer and a small, ovoid neuron in the suprapyramidal layer were investigated in the electron microscope. Some of these sections were processed for GABA immunogold cytochemistry. The soma and large dendrites of both neurons displayed GABA immunogold labelling. On the soma of medium-sized local circuit neuron there were numerous terminals; on the soma of the small one relatively fewer terminals were observed. The terminals contained round and/or flat synaptic vesicles. The long axonal branches of the neurons exhibited varicosities containing flattened or pleomorphic vesicles. Axo-dendritic, axo-somatic and a few axo-axonic synapses were observed. The large dense axon arborisation field of medium-sized local circuit neurons is properly situated to modulate intrinsic hippocampal activity and that of the small local circuit neurons is well situated to modulate the hippocampal input in the suprapyramidal layer.
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