Spiny Stellate Neurons Research Articles

Synaptic output of physiologically identified spiny stellate neurons in cat visual cortex.

Spiny stellate neurons of area 17 of the cat's visual cortex were physiologically characterised and injected intracellularly with horseradish peroxidase. Six neurons from sublamina 4A were selected. Five had the S-type of simple receptive fields; one had a complex receptive field. Their axons formed boutons mainly in layers 3 and 4. An electron microscopic examination of 45 boutons showed that each bouton formed one asymmetric synapse on average. Spines were the most frequent synaptic target (74%); dendritic shafts formed the remainder (26%). On the basis of ultrastructural characteristics, 8% of the target dendrites were characterised as originating from smooth gamma-aminobutyrate-ergic (GABAergic) neurons. Thus the major output of spiny stellate neurons is to other spiny neurons, probably pyramidal neurons in layer 3 and spiny stellates in layer 4.

Projection of the entorhinal layer II neurons in the rat as revealed by intracellular pressure-injection of neurobiotin.

A component of the perforant path, projection of the entorhinal layer II neurons, was investigated by recovering intracellularly labeled layer II neurons from the medial or intermediate entorhinal cortex in rats. The labeled layer II spiny stellate neurons had axon collaterals in layers I, II, and III of the entorhinal cortex as well as some axon collaterals in the subiculum. The stem axons gave rise to terminal axon branches that covered the entire extent (suprapyramidal blade, crest, and infrapyramidal blade) of the dentate gyrus and the CA2-3 fields in the transverse plane, forming a sheet-like formation. The axon arbor in the hippocampal formation spread up to 2 mm wide in a septotemporal direction. The sheet-like formation of the axon arbors was a narrow layer in the suprapyramidal blade and in the stratum lacunosum-moleculare of the CA2-3 fields. The layer became wider in the crest and infrapyramidal blade of the dentate gyrus. This study shows that the entorhinohippocampal circuit is not a simple circle from single cells level.

Form, function, and intracortical projections of neurons in the striate cortex of the monkey Macacus nemestrinus.

Single neurons were recorded in the striate visual cortex (area 17) of the old-world monkey Macacus nemestrinus. Eight pyramidal neurons, seven spiny stellate neurons, two basket cells, a clutch cell, and a chandelier cell were filled intracellularly with HRP. Their receptive fields were consistent with previous single-unit studies. Their axonal arbors were less elaborate than in equivalent neurons in the cat, but the laminar specificity of the boutons was very much more precise in the monkey than in the cat. Nevertheless, the basic cortical circuits in cat and monkey appear to be very similar.

Postnatal development of thalamic recipient neurons in the monkey striate cortex: Somatic inhibitory synapse acquisition by spiny stellate neurons of layer 4C

The development of type 2 (Colonnier, '81) synapses on the cell bodies of thalamic recipient spiny stellate neurons in layers 4C alpha and 4C beta of primary visual cortex neurons was examined over the first 36 postnatal weeks and in the adult monkey. The type 2 synapses, known to be GABAergic (Ribak, '78) and therefore presumed to be inhibitory, developed faster on the alpha neurons than the beta neurons. Both neuron groups show a marked increase and then decline in the percentage of the somatic membrane covered by type 2 synaptic appositions during this 36-week time period. The time course of the type 2 synapses development is compared to that of the spine synapse development described in previous studies (Lund and Holbach, '91; Lund et al., '91), and it is clear that on both neuron groups this inhibitory synapse population is put in place and refined later than the spine synapses. These findings suggest that each cortical neural circuit has a unique time course for its early development within an overall time window (Rakic et al., '86), or sensitive period (Hubel and Wiesel, '70). Visual deprivation, although causing the alpha and beta neurons to adopt a more similar temporal and numerical developmental pattern than normal, did not prevent acquisition and loss phases of type 2 synapses or the assumption of a normal numerical loading by 36 weeks of age.

Postnatal development of thalamic recipient neurons in the monkey striate cortex: Influence of afferent driving on spine acquisition and dendritic growth of layer 4c spiny stellate neurons

This study uses Golgi-impregnated material to examine the effects of altering the nature of afferent driving on the maturation of spines and dendrites on thalamic recipient spiny stellate neurons in layers 4C alpha and beta of the monkey striate cortex. These two laminae receive input from different sets of thalamic afferents with different functional properties. The development of dendritic spine and dendritic branch populations on these neurons in experimental animals is compared to the same features on similar groups of neurons in a series of normal animals described in the preceding study (Lund and Holbach, '91). Three conditions of rearing were used to alter afferent driving from normal: complete darkness (with in some cases return to normal diurnal light-dark cycle), bilateral eye lid suture, and monocular eye lid suture. Some of the normal and dark-reared infant monkeys were examined behaviorally for visual capacity in an earlier study (Regal et al., '76). All conditions of abnormal afferent driving caused changes from the normal developmental patterns of spine and dendritic arbor growth in these first-order neurons of the cortex and each condition differed in the nature of change produced. Major findings of this study are: 1. Vigorous spine acquisition and dendritic growth occurs under all conditions of visual deprivation on alpha and beta neurons. Eventual spine and dendritic attrition occurs under at least conditions of bilateral or monocular lid suture to produce a rather constant adult morphology. We assume, therefore, that visually driven activity is a modulator or shaper of the developmental process for thalamic recipient neurons of visual cortex, rather than being an initiator, terminator, or driving force for their maturation. 2. An innate "clock," whose nature is unknown but is apparently not driven by visual input, initiates and terminates a period of growth of the thalamic recipient neurons between birth and 30-32 weeks of age. 3. Factors controlling dendritic arbor growth and retraction are different from those controlling spine synapse addition or attrition. 4. Whereas the alpha and beta neurons normally show quite different early growth patterns between birth and 30 weeks of age, when both eyes are simultaneously deprived of vision, an early temporal and numerical convergence occurs in patterns of spine population development on the two groups of neurons. This convergent pattern assumes a different form in dark-reared and lid-sutured animals.(ABSTRACT TRUNCATED AT 400 WORDS)

Postnatal development of thalamic recipient neurons in the monkey striate cortex: Comparison of spine acquisition and dendritic growth of layer 4C alpha and beta spiny stellate neurons

A quantitative study has been made from Golgi impregnations of the maturation of dendrites and their spines on spiny stellate neurons in the macaque monkey primary visual cortex. The neurons studied lay within either the alpha or the beta division of lamina 4C; previous workers have shown the alpha division neurons to be contacted by thalamic axon terminals arising from the magnocellular division of the lateral geniculate nucleus (LGN) of the thalamus and the beta division neurons to be contacted by parvocellular LGN inputs. Most thalamic terminals and perhaps the majority of other type 1 (Colonnier, '81), presumed excitatory, inputs to these cells make synaptic contacts on the tips of their dendritic spines. Measurement was made of relative changes in the total number of spines on these alpha and beta spiny neurons over age by measuring both spine density along the dendrites and dendritic arbor size in single 90-microns sections from Golgi rapid preparations. Our previous work (Lund et al., '77; Boothe et al., '79) showed a marked proliferation and attrition of spines and dendritic branches to occur in the early postnatal weeks; Rakic et al. ('86) have since proposed that there is a cortexwide synchrony of synapse acquisition and loss during this same period. However, different visual capacities channelled via the magnocellular and parvicellular geniculate relays show different maturational rates (Harwerth et al., '86). This study indicates that the anatomical maturation of spines on the alpha and beta neurons is not temporally coincident from birth to 30 weeks. During this period, phases of spine acquisition and loss on alpha neurons precedes similar phases on beta neurons. The alpha neurons carry a peak spine population at 5-8 weeks postnatal, whereas the beta neurons carry their peak spine populations between 8 and 24 weeks postnatal. At all ages prior to 30 weeks, the two sets of neurons carry quite different total spine populations. Close to 30 weeks of age, the total spine coverage has fallen on both sets of neurons and becomes identical between the alpha and beta neurons. In animals aged 30 weeks to adult, spine coverage per neuron is maintained at a common figure for the alpha and beta neurons despite further growth and disparate dendritic arbor sizes and different local spine densities in the two groups; this suggests that some common sampling paradigm between pre- and postsynaptic elements is adopted by the alpha and beta neurons and also suggests the development of a close functional correlation between the two sets of neurons.

Cell surface molecules containing N-acetylgalactosamine are associated with basket cells and neurogliaform cells in cat visual cortex

In the cerebral cortex, the plant lectin Vicia villosa (VVA) selectively stains the surfaces of nonpyramidal neurons. This lectin binds specifically to alpha- and beta-linked N-acetylgalactosamine (GalNac). VVA-reactive carbohydrate is highly concentrated in layer 4 of the primary visual cortex of the cat, where it is associated exclusively with GABAergic local circuit neurons. We have studied this neuronal subset with intracellular electrophysiological recording and dye marking to identify the particular cell types expressing surface GalNac. Five different types of local circuit neurons were stained intracellularly (N = 45), but only 2 types, the columnar basket and large neurogliaform cells, were also labeled by the lectin (N = 19/45). Lectin negative types included small basket, chandelier, and large bitufted cells (26/45). Spiny stellate and pyramidal neurons were also lectin negative. Electrophysiological recordings revealed differences in the duration of action potentials in smooth versus spiny stellates but no differences between lectin-positive or -negative types. A biochemical analysis of cortical glycoproteins by SDS-PAGE and lectin blotting revealed multiple bands containing GalNac enriched in membrane fractions. These carbohydrate-containing molecules may be part of a biochemical mechanism for linking together cells with common functional properties.

Glutamate-positive neurons and axon terminals in cat sensory cortex: a correlative light and electron microscopic study.

Immunocytochemical methods were used to perform a correlative light and electron microscopic study of neurons and axon terminals immunoreactive to the antiglutamate (Glu) serum of Hepler et al. ('88) in the visual and somatic sensory areas of cats. At the light microscopic level, numerous Glu-positive neurons were found in all layers except layer I of both cortical areas. On the basis of the dendritic staining of Glu-positive cells, two major morphological categories were found: pyramidal cells, which were the most frequent type of immunostained neuron, and multipolar neurons, which were more numerous in layer IV of area 17 than in any other layer. A large number of Glu-positive neurons, however, did not display dendritic labelling and were considered unidentified neurons. Counts of labelled neurons were performed in the striate cortex; approximately 40% were Glu-positive. Numerous lightly stained punctate structures were observed in all cortical layers: the majority of these Glu-positive puncta were in the neuropil. After resectioning the plastic sections for electron microscopy it was observed that: 1) the majority of neurons unidentifiable at light microscopic level were indeed pyramidal neurons except in layer IV of area 17, where many stained cells were probably spiny stellate neurons. Some Glu-positive neurons, however, exhibited clear ultrastructural features of nonspiny nonpyramidal cells; 2) all synaptic contacts made by Glu-positive axon terminals were of the asymmetric type, but not all asymmetric synaptic contacts were labelled. The vast majority of postsynaptic targets of Glu-positive axons were unlabelled dendritic spines and shafts. The present results provide further evidence that Glu (or a closely related compound) is probably the neurotransmitter of numerous excitatory neurons in the neocortex.

The spiny stellate neurons in layer IV of the human auditory cortex. A Golgi study

The spiny stellate neurons have been studied by the Golgi method in the auditory koniocortex and parakoniocortex of man. Spiny stellate cells are a consistent though not very common component of layer IV. They are not confined to specific sublayers but occur at all depths of layer IV, and also in layer IIIc. Spiny stellate cells in the auditory areas show a great variety of their dendritic arborization pattern. The presence of all intermediate forms between small pyramidal cells—which constitute the dominant cell type in layer IV and which display an extraordinary heteromorphism—and spiny stellate cells shows the close kinship between both neuronal types. The morphology and distribution of spines along the dendrites of spiny stellate neurons are similar to those of the small pyramidal cells of the same layer. The axons, which were impregnated only in their proximal portions, mostly descend, giving rise to recurrent ascending collaterals, but initially ascending axons do also occur. Spiny stellate neurons are present in the different cytoarchitectonic areas examined, and thus they are not confined to the auditory koniocortex.

Structural alterations of dendritic spines induced by neural degeneration of their presynaptic afferents.

Morphological parameters were compared for dendritic spines of spiny stellate neurons in layer IV of the barrel region of mouse somatosensory cortex, which synapse with degenerated thalamocortical afferents (TC spines) and with intact, unidentified axon terminals (UI spines). Spiny stellate neurons were labeled for light and electron microscopic identification by Golgi impregnation and gold toning. Dendritic spines were examined in series of thin sections, and TC spines were ultrastructurally detectable because of the degeneration-induced characteristic appearance of the TC axon terminals. Results show that the means of the width of the spine head and of the length of the spine stalk were significantly higher in TC spines than in UI spines by about 11 and 25%, respectively. The variability of these two morphological parameters was significantly lower for TC spines. The mean of the spine stalk width at the narrowest cross section of the stalk was about 0.12 microns, with no significant difference observed between the two spine groups. No specific relationship was found in either the TC or the UI groups of spines between the length of the spine stalk and the width of the spine stalk at its narrowest profile. As structural features typifying transneuronal degeneration were not observed along the dendritic spines examined, it is speculated that the morphological differences encountered between the TC and UI spines may result, at least in part, from the degeneration-induced synaptic inactivity of the TC axospinous synapses, rather than exclusively from any direct effects of the degeneration process.

Quantification of thalamocortical synapses with spiny stellate neurons in layer IV of mouse somatosensory cortex.

The distribution of thalamocortical (TC) and other synapses involving spiny stellate neurons in layer IV of the barrel region of mouse primary somatosensory cortex (SmI) was examined in seven male CD/1 mice. TC axon terminals were labeled by lesion-induced degeneration, which has been shown to label reliably all TC synapses in mouse barrel cortex. Spiny stellate neurons, labeled by Golgi impregnation and gold toning, were identified with the light microscope prior to thin sectioning and electron microscopy. Analysis of eight dendritic segments from seven spiny stellate neurons showed that most of their synapses are with their dendritic spines, rather than with their shafts. Axospinous synapses are primarily of the asymmetrical type, whereas axodendritic synapses are mainly of the symmetrical type. Dendrites of spiny stellate neurons consistently form thalamocortical synapses, most of which involve spine heads rather than spine stalks or dendritic shafts. From 10.4% to 22.9% of all asymmetrical synapses with dendrites of spiny stellate neurons involve TC axon terminals. In general, this is a higher range than the ranges that characterize the TC synaptic connectivity of dendrites belonging to other types of neurons, implying that spiny stellate neurons are perhaps more strongly influenced by TC synaptic input than other types of cortical neurons examined previously. Spines involved in TC synapses were distributed irregularly along each of the stellate cell dendrites; about half of the interspinous intervals between these spines were about 5 microns or less. Modulations of the efficacy of TC synaptic input to dendrites of layer IV spiny stellate neurons are discussed in the light of recently reported computer simulated analyses of axospinous synaptic connections.

Synaptic relationships of a type of GABA-immunoreactive neuron (clutch cell), spiny stellate cells and lateral geniculate nucleus afferents in layer IVC of the monkey striate cortex

The precise stimulus specificity of striate cortical neurons is strongly influenced by processes involving gamma-aminobutyric acid (GABA). In the visual cortex of the monkey most afferents from the lateral geniculate nucleus terminate in layer IVC. We identified a type of smooth dendritic neuron (clutch cell) that was immunoreactive for GABA, and whose Golgi-impregnated dendrites and axon were largely restricted to layer IVCβ. The slightly ovoid somata were 8–12μm by 12–15μm and the dendritic field was often elongated, extending 80–200 μm in one dimension. The axonal field was 100–150 μm in diameter and densely packed with large bulbous boutons. Although mainly located in IVCβ both the dendritic and axonal processes entered IVCα. Fine structural features of GABA-immunoreactive and-impregnated clutch cells and impregnated spiny stellate cells were compared. Clutch cells had more cytoplasm, more densely packed mitochondria and endoplasmic reticulum, and made type II as opposed to type I synapses. A random sample of 159 elements postsynaptic to three clutch cells showed that they mainly terminated on dendritic shafts (43.8–58.5%) and spines (20.8–46.3%), rather than somata (10–17%). The majority of the postsynaptic targets were characteristic of spiny stellate cells. This was directly demonstrated by studying synaptic contacts between an identified GABA positive clutch cell and the dendrites and soma of an identified spiny stellate cell. The termination of clutch cells mainly on dendrites and spines of spiny stellate cells suggests that they interact with other inputs to the same cells. Following an electrolytic lesion in the ipsilateral lateral geniculate nucleus we examined the distribution of degenerating terminals on three identified spiny stellate neurons in layer IVCβ. Out of eight synapses from the lateral geniculate nucleus one was on a dendritic shaft, the rest on spines. Only a small fraction of all synapses on the cells were from degenerating boutons. A clutch cell within the area of dense terminal degeneration was not contacted by terminals from the lateral geniculate nucleus. The results show that layer IVC in the monkey has a specialized GABAergic neuron that terminates on spiny stellate cells monosynaptically innervated from the lateral geniculate nucleus. Possible functions of clutch cells may include inhibitory gating of geniculate input to cortex; maintenance of the antagonistic subregions in the receptive fields; and the creation from single opponent of double colour opponent receptive fields.

Stability of dendrites in cortical barrels of C57BL/6N mice between 4 and 45 months

Qualitative and quantitative studies of dendritic parameters were conducted on Golgi-impregnated layer IV spiny stellate neurons in the posteromedial barrel subfield (PMBSF) of somatosensory cortex of the C57B1/6N mouse. Three mice each, at 4, 12, 22, 26, 30, 36 and 45 months of age were studied. No qualitative changes were observed among animals of different ages. The quantitative data indicated that dendritic length and numbers of segments remained unchanged over all ages studied.

Synaptic connections of intracellularly filled clutch cells: a type of small basket cell in the visual cortex of the cat.

Light and electron microscopic quantitative analysis was carried out on a type of neuron intracellularly filled with horseradish peroxidase. Two cells were studied in area 17, one of which was injected intra-axonally, and its soma was not recovered. One cell was studied in area 18. The two somata were on the border of layers IVa/b; they were radially elongated and received synapses from numerous large boutons with round synaptic vesicles. The dendrites were smooth and remained largely in layer IV. The cells can be recognised on the basis of their axonal arbor, which was restricted to layer IV (90-95% of boutons) with minor projections to layers III, V, and VI. Many of the large, bulbous boutons contacted neuronal somata, short collaterals often forming "claw"-like configurations around cells. The name "clutch cell" is suggested to delineate this type of neuron from other aspiny multipolar cells. Computer-assisted reconstruction of the axon showed that in layer IV the axons occupied a rectangular area about 300 X 500 microns, elongated anteroposteriorly in area 17 and mediolaterally in area 18. The distributions of synaptic boutons and postsynaptic cells were patchy within this area. A total of 321 boutons were serially sectioned in area 17. The boutons formed type II synaptic contacts. The postsynaptic targets were somata (20-30%), dendritic shafts (35-50%), spines (30%), and rarely axon initial segments. Most of the postsynaptic somata tested were not immunoreactive for GABA and their fine structural features suggest that they are spiny stellate, star pyramidal, and pyramidal neurons. The characteristics of most of the postsynaptic dendrites and spines also suggest that they belong to these spiny neurons. A few of the postsynaptic dendrites and somata exhibited characteristics of cells with smooth dendrites and these somata were immunoreactive for GABA. It is suggested that clutch cells are inhibitory interneurons exerting their effect mainly on layer IV spiny neurons in an area localised perhaps to a single ocular dominance column. The specific laminar location of the axons of clutch cell also suggests that they may be associated with the afferent terminals of lateral geniculate nucleus cells, and could thus be responsible for generating some of the selective properties of neurons of the first stage of cortical processing.

The morphology and synaptic connections of spiny stellate neurons in monkey visual cortex (area 17): a Golgi-electron microscopic study.

Based on a gold-toning, Golgi-electron microscope examination of 12 small and medium-sized spiny stellate neurons in laminae 4A, 4B, and 4C of the monkey visual cortex (area 17), the ultrastructure of the cell somata, dendrites, and axons of these neurons is described. Particular attention is paid to the synapses involving the surface of different parts of these neurons. Only symmetric synapses occur on the somata of spiny stellate neurons, and these occur with a frequency of 11.0-15.9 synapses/100 microns2 perikaryal surface. Symmetric synapses also occur on dendritic shafts and, occasionally, on dendritic spines. Asymmetric synapses are occasionally present along the dendritic shafts of spiny stellate neurons, but the majority of asymmetric synapses (75-95%) occur on their dendritic spines. The initial axon segments of the smallest spiny stellate neurons possess no axo-axonal synapses, but several symmetric synapses are present along the initial segment of a medium-sized, spiny stellate neuron in layer 4B. Fifty-three synapses made by boutons of the axons of these spiny stellate neurons have been identified, and all are asymmetric. Sixty per cent of the synapses are formed by boutons en passant and the remainder by the terminal swellings of spine-like axonal appendages, boutons terminaux. Of the synapses formed by the axons of spiny stellate cells, axo-spinous synapses outnumber axo-dendritic synapses two to one, and axo-dendritic synapses involve both spinous and aspinous dendrites. Evidence is presented which suggests that many of the axon terminals forming asymmetric synapses with the dendritic shafts and spines of spiny stellate neurons are derived from other spiny stellate neurons.

Characterization by Golgi impregnation of neurons that accumulate 3H-GABA in the visual cortex of monkey.

3H-GABA was injected into restricted regions of visual areas 1 and 2 (cortical areas 17 and 18) on the lateral surface of the occipital lobe in monkeys. The injected tissue was processed for Golgi impregnation and gold toning. Sections containing Golgi-impregnated neurons were re-embedded, sectioned at 1 micron, and prepared for autoradiography to reveal neurones that had selectively accumulated 3H-GABA. Golgi-impregnated pyramidal, spiny stellate and aspiny nonpyramidal neurons were examined for 3H-GABA accumulation. Out of 47 aspiny non-pyramidal neurons 16 were labelled by 3H-GABA. The other cell types did not accumulate the amino acid. Twelve of the labelled neurons were drawn. Eight were bitufted neurons with their dendrites oriented predominantly radially, three were small multipolar neurons, and one could be reconstructed only partially. One neuron had a locally arborizing axon in layer III. Two bitufted, Golgi-impregnated neurons in layer II and upper III of area 18 were labelled from GABA injection radially beneath in layer VI, providing evidence for earlier suggestions that in the monkey's visual cortex the cells in the upper layers which project radially and accumulate 3H-GABA are aspiny non-pyramidal cells. The results indicate the existence of different types of putative GABA-ergic interneurons.

Spine formation and maturation of type 1 synapses on spiny stellate neurons in primate visual cortex

This study continues an investigation (Mates and Lund, '83a) of neuronal development in lamina 4C of macaque monkey striate visual cortex. The maturational history of the type 1 synaptic contacts on spine-bearing stellate neurons, which comprise 95% of the neurons of the lamina, is described. It is shown that type 1 contacts are initially found on the dendritic shafts; these contacts appear to be carried out on spine outgrowths. This leads to the adult condition where type 1 contacts are found only on the spine tips. A similar phenomenon has been reported for pyramidal neurons of the rat (reported during the course of this study by Miller and Peters, '81). In later maturation the spine and its type 1 contact may be lost; profiles found in the neuropil illustrate a process of shrinkage and detachment of both the type 1 axon terminals and postsynaptic dendritic spines in normal maturation. These findings provide an explanation for the marked increase and decrease in spine populations observed to occur on these neurons during normal maturation in an earlier study by Boothe et al. ('79).

Developmental changes in the relationship between type 2 synapses and spiny neurons in the monkey visual cortex.

This study continues an exploration of synaptic development in the primary visual cortex of the monkey (Macaca nemestrina). In a prior study (Mates and Lund, '83a), we observed that type 2 synapses on the cell bodies of spiny stellate neurons of lamina 4C appeared not only to increase in number during early postnatal development but also subsequently decreased during maturation. Using quantitative, stereological electron microscopic methods, we examined the maturation of this synapse population from embryonic day 159 to adult, on spiny stellate neurons of 4C alpha and beta and, for comparison, on pyramidal neurons in upper and lower lamina 6. Tissue was also taken for comparison from two animals reared to 8 weeks of age with binocular eyelid closure from birth. We confirmed that a marked increase and subsequent decrease occurred in this somal type 2 synapse population on both neuron populations. However, due to the infrequency of the smooth dendritic neurons (approximately 5% of the neuron population) giving rise to the type 2 contacts, and due to expansion of the neuropil during maturation increasing intercell distances against constant volume of the type 2 axon arbors, it is concluded that the decrease in type 2 somal synapses may represent a redistribution to dendrites rather than loss from the neuropil. Cells of lamina 4C beta (receiving input from the parvocellular lateral geniculate nucleus-LGN) show a slower initial accumulation of type 2 contacts compared to neurons of lamina 4C alpha (receiving input from magnocellular LGN), or to pyramidal neurons of lamina 6.(ABSTRACT TRUNCATED AT 250 WORDS)

A quantitative investigation of spine and dendrite development of neurons in visual cortex (area 17) of Macaca nemestrina monkeys.

In a previous Golgi study (Lund et al., '77) which examined the development of the macaque monkey striate cortex (area 17) it was observed that the dendrites of neurons within the visual cortex show a marked increase in the number of spines on their surface during the first eight weeks of postnatal life. The qualitative observation was also made that all neurons then showed a marked decrease in spine numbers by the time the animal was adult. Since these spines are known to be sites of synaptic contact, changes in their numbers may reflect changes in synapse populations on these neurons. This study examines quantitatively spine frequency and total dendritic development of Golgi impregnated neurons in monkeys ranging in age from 145 days gestation to adult. Four cell types were studied: spiny stellate neurons from laminae IVCalpha and IVCbeta and pyramidal neurons with soma in either lamina IIIB or upper lamina VI. After consideration of possible sources of variation in spine numbers several conclusions are made: (1) Dendritic spine development appears to be a tightly controlled process both in terms of actual numbers of spines on a neuron at any one age and in the rate of change of spine frequency. (2) The neurons populations examined all show a gradual increase in spine numbers up to eight weeks of age. (3) At least two different trends are found in spine population maturation after the eight week point: (A)-the spine population may remain constant at the eight week level for same period of time or (B)-there may be a rapid decline in spine numbers following the eight week peak. (4) There is a suggestion that those neurons associated with direct input, or early stages in the relays, from the parvocellular geniculate laminae show trend B, while those associated with magnocellular input, or later order combined relays within the cortex, show trend A. (5) Different parts of a single pyramidal neuron dendrite may show either trend A or trend B, depending on the lamina location of the dendritic segment considered. (6) All neurons show spine population decreases between nine months of age and adult (5-7 years) suggesting continuing long term maturational changes.

Anatomical organization of the primary visual cortex (area 17) of the cat. A comparison with area 17 of the macaque monkey

Golgi and axonal transport techniques have been used to examine the organization of neurons within primary visual cortex, area 17, of the cat. This organization has been compared to that of the primate cortical area 17 as described in previous studies and it is discussed in relationship to the distribution of afferents from the dorsal lateral geniculate nucleus (dLGN). The visual cortex of the cat and monkey show strong similarities in the laminar positions of neurons projecting extrinsically and also in the restriction of spiny stellate neurons to a central lamina (lamina 4) receiving input from the dLGN. However, lamina 4B in the monkey, which contains spiny stellate neurons but does not receive direct input from the dLGN, has no direct counterpart in cat area 17. Axon projections of spiny stellate neurons in the other divisions of lamina 4 differ in cat and monkey: the small, closely packed neurons in the lowermost division of lamina 4 (4B in the cat, 4Cbeta in the monkey) project chiefly within lamina 4 in the cat whereas in the monkey they have a strong projection to lamina 3. In the cat, spiny stellate neurons of lamina 4A project upon lamina 3 whereas in the monkey those in the apparently equivalent zone, 4Calpha, project upon lamina 4B. Most non-spiny stellate neurons examined have precisely organized interlaminar axonal projections which differ from the axon trajectories of neighboring spiny neurons.