scholarly journals Excitatory inputs to spiny cells in layers 4 and 6 of cat striate cortex

2002 ◽  
Vol 357 (1428) ◽  
pp. 1793-1808 ◽  
Author(s):  
N.J. Bannister ◽  
J.C. Nelson ◽  
J.J.B. Jack

The principal target of lateral geniculate nucleus in the cat visual cortex is the stellate neurons of layer 4. In previously reported work with intracellular recording and extracellular stimulation in slices of visual cortex, three general classes of fast excitatory synaptic potentials (EPSPs) in layer 4a spiny stellate neurons were identified. One of these classes, characterized by large and relatively invariant amplitudes (mean 1.7 mV, average coefficient of variation (CV) 0.083) were attributed to the action of geniculate axons because, unlike the other two classes, they could not be matched by intracortical inputs, using paired recording. We have examined in detail the properties of this synaptic input in twelve examples, selecting for study those EPSPs where there was secure extracellular stimulation of the single fibre input to a pair of stimuli 50 ms apart. In our analysis, we conclude that the depression that these inputs show to the second stimulus is entirely postsynaptic, since the evidence strongly suggests that the probability of transmitter release at the synaptic site(s) remains 1.0 for both stimuli. We argue that the most plausible explanation for this postsynaptic depression is a reduction in the average probability of opening the synaptic channels. Using a simple biochemical analysis (c.f. Sigworth plot), it is then possible to calculate the number of synaptic channels and their probability of opening, for each of the 12 connections. The EPSPs had a mean amplitude of 1.91 mV (±1.3 mV SD) and a mean CV of 0.067 (± 0.022). The calculated number of channels ranged from 20 to 158 (59.4 ± 48.7) and their probability of opening to the first EPSP had an average of 0.83 (± 0.09), with an average depression of the probability to 0.60 for the second EPSP. Geniculate afferents also terminate in layer 6. Intracellular recordings were also made in the upper part of this layer and a total of 51 EPSPs were recorded from pyramidal cells of three principal types. Amongst this dataset we sought EPSPs with similar properties to those characterized in layer 4a. Three examples were found, which is a much lower percentage (6%) than the incidence of putative geniculate EPSPs found in layer 4a (42%).

1990 ◽  
Vol 5 (1) ◽  
pp. 67-81 ◽  
Author(s):  
Edgar A. Deyoe ◽  
Susan Hockfield ◽  
Hideki Garren ◽  
David C. Van Essen

AbstractWe have examined the distribution of immunoreactivity for the monoclonal antibody Cat-301 in visual cortex of the macaque monkey. Remarkably, those portions of striate cortex (V1) and extrastriate cortex that are most immunoreactive for Cat-301 are anatomically interconnected and are dominated by inputs arising from the magnocellular layers of the LGN (which are themselves highly immunoreactive). In particular, we found that a band of Cat-301 labeled neurons known to exist in layer 4 of V1 is centered on the boundary between layers 4Cα and 4B and thus includes portions of both the primary target of the magnocellular LGN and its subsequent relay through layer 4B. We also demonstrated consistently strong Cat-301 immunoreactivity in all three extrastriate targets of layer 4B: areas V3, MT, and the cytochrome-oxidase (CO) enriched thick stripes of V2. In V2, there was a close correlation between Cat-301 labeling and clusters of cells projecting to MT but not to V4. This was true even in regions where the CO pattern was equivocal or irregular, indicating that Cat-301 is a more reliable marker than CO for the thick-stripe subregions of V2. Finally, we found strong Cat-301 immunoreactivity in at least parts of areas V3A, the MST complex, and the posterior parietal complex, but not in area V4 or inferotemporal cortex. The molecular specificity revealed by this single marker thus correlates with functionally specific subdivisions at each hierarchical level over nearly the entire known extent of the visual pathway in macaques. This supports the notion that these subdivisions form an anatomically, physiologically, and now molecularly distinct pathway known as the M-stream.


After injections of the enzyme horseradish peroxidase (HRP) into the superior colliculus of macaque monkeys, labelled cells in the neocortex were found to be restricted to layer V in all areas except striate visual cortex. In striate visual cortex, cortico-tectal cells were found both in layer V and in layer VI. The labelled cells in the two layers belonged to morphologically different populations: those of layer V were the common pyramidal cells and those of layer VI were identified as solitary cells of Meynert. This finding may provide new insights into the physiology of the cortico-collicular pathways. It also shows that the striate area in primates differs, with respect to cortico-tectal laminar specificity, from other neocortex.


2021 ◽  
Vol 11 (1) ◽  
Author(s):  
Rongkang Deng ◽  
Joseph P. Y. Kao ◽  
Patrick O. Kanold

AbstractThe development of GABAergic interneurons is important for the functional maturation of cortical circuits. After migrating into the cortex, GABAergic interneurons start to receive glutamatergic connections from cortical excitatory neurons and thus gradually become integrated into cortical circuits. These glutamatergic connections are mediated by glutamate receptors including AMPA and NMDA receptors and the ratio of AMPA to NMDA receptors decreases during development. Since previous studies have shown that retinal input can regulate the early development of connections along the visual pathway, we investigated if the maturation of glutamatergic inputs to GABAergic interneurons in the visual cortex requires retinal input. We mapped the spatial pattern of glutamatergic connections to layer 4 (L4) GABAergic interneurons in mouse visual cortex at around postnatal day (P) 16 by laser-scanning photostimulation and investigated the effect of binocular enucleations at P1/P2 on these patterns. Gad2-positive interneurons in enucleated animals showed an increased fraction of AMPAR-mediated input from L2/3 and a decreased fraction of input from L5/6. Parvalbumin-expressing (PV) interneurons showed similar changes in relative connectivity. NMDAR-only input was largely unchanged by enucleation. Our results show that retinal input sculpts the integration of interneurons into V1 circuits and suggest that the development of AMPAR- and NMDAR-only connections might be regulated differently.


2013 ◽  
Vol 33 (28) ◽  
pp. 11372-11389 ◽  
Author(s):  
J. Zhuang ◽  
C. R. Stoelzel ◽  
Y. Bereshpolova ◽  
J. M. Huff ◽  
X. Hei ◽  
...  

Nature ◽  
1986 ◽  
Vol 320 (6060) ◽  
pp. 310-311 ◽  
Author(s):  
Simon LeVay
Keyword(s):  

1991 ◽  
Vol 3 (1) ◽  
pp. 44-53 ◽  
Author(s):  
Sidney R. Lehky ◽  
Randolph Blake

It is proposed that inputs to binocular cells are gated by reciprocal inhibition between neurons located either in the lateral geniculate nucleus or in layer 4 of striate cortex. The strength of inhibitory coupling in the gating circuitry is modulated by layer 6 neurons, which are the outputs of binocular matching circuitry. If binocular inputs are matched, the inhibition is modulated to be weak, leading to fused vision, whereas if the binocular inputs are unmatched, inhibition is modulated to be strong, leading to rivalrous oscillations. These proposals are buttressed by psychophysical experiments measuring the strength of adaptational aftereffects following exposure to an adapting stimulus visible only intermittently during binocular rivalry.


Of the many possible functions of the macaque monkey primary visual cortex (striate cortex, area 17) two are now fairly well understood. First, the incoming information from the lateral geniculate bodies is rearranged so that most cells in the striate cortex respond to specifically oriented line segments, and, second, information originating from the two eyes converges upon single cells. The rearrangement and convergence do not take place immediately, however: in layer IVc, where the bulk of the afferents terminate, virtually all cells have fields with circular symmetry and are strictly monocular, driven from the left eye or from the right, but not both; at subsequent stages, in layers above and below IVc, most cells show orientation specificity, and about half are binocular. In a binocular cell the receptive fields in the two eyes are on corresponding regions in the two retinas and are identical in structure, but one eye is usually more effective than the other in influencing the cell; all shades of ocular dominance are seen. These two functions are strongly reflected in the architecture of the cortex, in that cells with common physiological properties are grouped together in vertically organized systems of columns. In an ocular dominance column all cells respond preferentially to the same eye. By four independent anatomical methods it has been shown that these columns have the form of vertically disposed alternating left-eye and right-eye slabs, which in horizontal section form alternating stripes about 400 μm thick, with occasional bifurcations and blind endings. Cells of like orientation specificity are known from physiological recordings to be similarly grouped in much narrower vertical sheeet-like aggregations, stacked in orderly sequences so that on traversing the cortex tangentially one normally encounters a succession of small shifts in orientation, clockwise or counterclockwise; a 1 mm traverse is usually accompanied by one or several full rotations through 180°, broken at times by reversals in direction of rotation and occasionally by large abrupt shifts. A full complement of columns, of either type, left-plus-right eye or a complete 180° sequence, is termed a hypercolumn. Columns (and hence hypercolumns) have roughly the same width throughout the binocular part of the cortex. The two independent systems of hypercolumns are engrafted upon the well known topographic representation of the visual field. The receptive fields mapped in a vertical penetration through cortex show a scatter in position roughly equal to the average size of the fields themselves, and the area thus covered, the aggregate receptive field, increases with distance from the fovea. A parallel increase is seen in reciprocal magnification (the number of degrees of visual field corresponding to 1 mm of cortex). Over most or all of the striate cortex a movement of 1-2 mm, traversing several hypercolumns, is accompanied by a movement through the visual field about equal in size to the local aggregate receptive field. Thus any 1-2 mm block of cortex contains roughly the machinery needed to subserve an aggregate receptive field. In the cortex the fall-off in detail with which the visual field is analysed, as one moves out from the foveal area, is accompanied not by a reduction in thickness of layers, as is found in the retina, but by a reduction in the area of cortex (and hence the number of columnar units) devoted to a given amount of visual field: unlike the retina, the striate cortex is virtually uniform morphologically but varies in magnification. In most respects the above description fits the newborn monkey just as well as the adult, suggesting that area 17 is largely genetically programmed. The ocular dominance columns, however, are not fully developed at birth, since the geniculate terminals belonging to one eye occupy layer IVc throughout its length, segregating out into separate columns only after about the first 6 weeks, whether or not the animal has visual experience. If one eye is sutured closed during this early period the columns belonging to that eye become shrunken and their companions correspondingly expanded. This would seem to be at least in part the result of interference with normal maturation, though sprouting and retraction of axon terminals are not excluded.


1997 ◽  
Vol 77 (4) ◽  
pp. 1939-1949 ◽  
Author(s):  
Mohamed Ouardouz ◽  
Jean-Claude Lacaille

Ouardouz, Mohamed and Jean-Claude Lacaille. Properties of unitary IPSCs in hippocampal pyramidal cells originating from different types of interneurons in young rats. J. Neurophysiol. 77: 1939–1949, 1997. Whole cell recordings were used in hippocampal slices of young rats to examine unitary inhibitory postsynaptic currents (uIPSCs) evoked in CA1 pyramidal cells at room temperature. Loose cell-attached stimulation was applied to activate single interneurons of different subtypes located in stratum oriens (OR), near stratum pyramidale (PYR), and at the border of stratum radiatum and lacunosum-moleculare (LM). uIPSCs evoked by stimulation of PYR and OR interneurons had similar onset latency, rise time, peak amplitude, and decay. In contrast, uIPSCs elicited by activation of LM interneurons were significantly smaller in amplitude and had a slower time course. The mean reversal potential of uIPSCs was −53.1 ± 2.1 (SE) mV during recordings with intracellular solution containing potassium gluconate. With the use of recording solution containing the potassium channel blocker cesium, the reversal potential of uIPSCs was not significantly different (−58.5 ± 2.6 mV), suggesting that these synaptic currents were not mediated by potassium conductances. Bath application of the γ-aminobutyric acid-A (GABAA) receptor antagonist bicuculline (25 μM) reversibly blocked uIPSCs evoked by stimulation of all interneuron subtypes. In bicuculline, the mean peak amplitude of uIPSCs recorded with potassium gluconate was reduced to 3.5 ± 4.4% of control ( n = 7). Similarly, with cesium methanesulfonate, the mean amplitude in bicuculline was 2.9 ± 3.1% of control ( n = 13). Application of the GABAB receptor antagonist CGP 55845A (5 μM) resulted in a significant and reversible increase in the mean amplitude of uIPSCs recorded with cesium-containing intracellular solution. Thus uIPSCs from all cell types appeared under tonic presynaptic inhibition by GABAB receptors. Paired stimulation of individual interneurons at 100- to 200-ms intervals did not result in paired pulse depression of uIPSCs. For individual responses, a significant negative correlation was observed between the amplitude of the first and second uIPSCs. A significant paired pulse facilitation (154.0 ± 8.0%) was observed when the first uIPSC was smaller than the mean of all first uIPSCs. A small, but not significant, paired pulse depression (90.8 ± 4.0%) was found when the first uIPSC was larger than the mean of all first uIPSCs. Our results indicate that these different subtypes of hippocampal interneurons generate Cl−-mediated GABAA uIPSCs. uIPSCs originating from different types of interneurons may have heterogeneous properties and may be subject to tonic presynaptic inhibition via heterosynaptic GABAB receptors. These results suggest a specialization of function for inhibitory interneurons and point to complex presynaptic modulation of interneuron function.


1997 ◽  
Vol 77 (6) ◽  
pp. 2879-2909 ◽  
Author(s):  
Izumi Ohzawa ◽  
Gregory C. Deangelis ◽  
Ralph D. Freeman

Ohzawa, Izumi, Gregory C. DeAngelis, and Ralph D. Freeman. Encoding of binocular disparity by complex cells in the cat's visual cortex. J. Neurophysiol. 77: 2879–2909, 1997. To examine the roles that complex cells play in stereopsis, we have recorded extracellularly from isolated single neurons in the striate cortex of anesthetized paralyzed cats. We measured binocular responses of complex cells using a comprehensive stimulus set that encompasses all possible combinations of positions over the receptive fields for the two eyes. For a given position combination, stimulus contrast could be the same for the two eyes (2 bright or 2 dark bars) or opposite (1 bright and 1 dark). These measurements provide a binocular receptive field (RF) profile that completely characterizes complex cell responses in a joint domain of left and right stimulus positions. Complex cells typically exhibit a strong selectivity for binocular disparity, but are only broadly selective for stimulus position. For most cells, selectivity for disparity is more than twice as narrow as that for position. These characteristics are highly desirable if we assume that a disparity sensor should exhibit position invariance while encoding small changes in stimulus depth. Complex cells have nearly identical binocular RFs for bright and dark stimuli as long as the sign of stimulus contrast is the same for the two eyes. When stimulus contrast is opposite, the binocular RF also is inverted such that excitatory subregions become suppressive. We have developed a disparity energy model that accounts for the behavior of disparity-sensitive complex cells. This is a hierarchical model that incorporates specific constraints on the selection of simple cells from which a complex cell receives input. Experimental data are used to examine quantitatively predictions of the model. Responses of complex cells generally agree well with predictions of the disparity energy model. However, various types of deviations from the predictions also are found, including a highly elongated excitatory region beyond that supported by a single energy mechanism. Complex cells in the visual cortex appear to provide a next level of abstraction in encoding information for stereopsis based on the activity of a group of simple-type subunits. In addition to exhibiting narrow disparity tuning and position invariance, these cells seem to provide a partial solution to the stereo correspondence problem that arises in complex natural scenes. Based on their binocular response properties, these cells provide a substantial reduction in the complexity of the correspondence problem.


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