The Functional Organization of Local Circuits in Visual Cortex: Insights from the Study of Tree Shrew Striate Cortex

We have used a combination of anatomical and physiological techniquesto explore the functional organization of vertical and horizontalconnections in tree shrew striate cortex. Our studies of verticalconnections reveal a remarkable specificity in the laminar arrangementof the projections from layer IV to layer III that establishesthree parallel intracortical pathways. The pathways that emergefrom layer IV are not simple continuations of parallel thalamocorticalpathways. Layer IV and its connections with layer II/III restructurethe inputs from the LGN, combining the activity from ON andOFF channels and from the left and right eye and transmit theproducts of this synthesis to separate strata within the overlyinglayers. In addition, studies of two other prominent verticalconnection pathways, the projections from layer VI to layerIV and from layer II/III to layer V suggest that the parallelnature of these systems is perpetuated throughout the corticaldepth. Our studies of horizontal connections have revealed a systematicrelationship between a neuron's orientation preference and thedistribution of its axon arbor across the cortical map of visualspace. Horizontal connections in layer II/III extend for greaterdistances and give rise to a greater number of terminals alongan axis of the visual field map that corresponds to the neuron'spreferred orientation. These findings suggest that the contributionof horizontal inputs to the response properties of layer II/IIIneurons is likely to be greater in regions of visual space thatlie along the axis of preferred orientation (endzones) thanalong the orthogonal axis (side zones). Topographically alignedhorizontal connections may contribute to the orientation preferenceof layer II/III neurons and could account for the axial specificityof some receptive field surround effects. Together, these results emphasize that specificity in the spatialarrangement of local circuit axon arbors plays an importantrole in shaping the response properties of neurons in visualcortex.     

Neuronal Organization in Area 17 of Cat Visual Cortex

An antibody to microtubule-associated protein 2 (MAP2) has beenused to examine the arrangements of neurons in striate cortexof the cat. It is found that the apical dendrites of medium-sizedand large pyramidal cells in layer V group together to formclusters that have a center-to-center spacing of about 56 µm.As these clusters ascend, the apical dendrites of pyramidalcells in layer ll/lll are added to them. The thinner apicaldendrites of the smaller pyramidal cells in layer VIa also formgroups that are referred to as bundles. These bundles ascendinto layer IV independent of the clusters, and their arrangementsuggests that the bundles are formed so that the apical dendritesof the layer VIa pyramids can pass between the groups of cellbodies of the layer V neurons. It is proposed that the clusters formed from the apical dendritesof the layer V and layer ll/lll pyramidal cells represent theaxes of vertical modules of pyramidal cells, which representthe basic neuronal aggregates within area 17 of cat visual cortex.And it is suggested that these modules can be recruited in variouscombinations, most obviously by the excitation provided by thethalamic inputs, to form the functional columns, such as theones that are concerned with eye preference and orientation.Based upon the distribution of the dendritic clusters, the pyramidalcell modules would have diameters of 56 µm, and sincethey are considered to extend through the depth of the cortex,each one would contain some 203 neurons. The striate cortexof one hemisphere contains 160, 000 of these modules, whichis about the same as the number of X-cells projecting to onehemisphere from the dorsal lateral geniculate nucleus and twicethe number of X- or ß-ganglion cells in the retina.The form of the pyramidal cell modules in cat striate cortexis compared to those present in monkey striate cortex, in whichthe similar modules are about 10 times more numerous, but only31 µm in diameter (Peters and Sethares, 1991a).     

Intracortical excitation of spiny neurons in layer 4 of cat striate cortex in vitro

Recordings were made from pairs of neurons in cat striate visual cortex in vitro to study the AMPA-channel-mediated components of intracortical excitatory synaptic connections between layer 4 spiny neurons and between layer 6 and layer 4 spiny neurons. Forty-six of the 72 cells recorded were identified morphologically. They consisted of spiny stellate and pyramidal cells in layer 4, and pyramidal cells in layer 6. Connections between layer 4 excitatory cells involve excitatory postsynaptic potentials (EPSPs) averaging 949 microV, with an average coefficient of variation of 0.21 (n = 30). The synapses operate at very high release probabilities (0.69-0.98). With repetitive stimulation these EPSPs show varying degrees of depression, largely mediated by presynaptic changes in release probability. Four pairs of layer 4 cells were reciprocally connected. The connections from layer 6 to layer 4 involve smaller, more variable EPSPs, with an average amplitude of 214 microV, and average coefficient of variation 0.72 (n = 7). These synapses operate at moderately high release probabilities (0.37-0.56). They show facilitation with repetitive stimulation, mediated largely by presynaptic changes in release probability. One excitatory connection from a layer 4 neuron to a layer 6 pyramidal cell was also detected. Thus, layer 4 spiny neurons receive effective excitation from two intracortical sources that have different synaptic dynamics and are likely to contribute significantly to the temporal properties of these cells in vivo.     

The occipitoparietal pathway of the macaque monkey: comparison of pyramidal cell morphology in layer III of functionally related cortical visual areas

The dendritic morphology of pyramidal cells located at the base of layer III in the primary visual area (V1), the second visual area (V2), the middle temporal area (MT), the ventral portion of the lateral intraparietal area (LIPv) and in the portion of cytoarchitectonic area 7a within the anterior bank of the superior temporal sulcus was revealed by injecting neurons with Lucifer Yellow in fixed, flattened slices of macaque monkey visual cortex. These areas correspond to different levels of the occipitoparietal cortical 'stream', which processes information related to motion and spatial relationships in the visual field. The tissue was immunocytochemically processed to obtain a light-stable diaminobenzidine reaction product, revealing the dendritic morphology in fine detail. Retrogradely labelled MT- projecting neurons in supragranular V1 (layer IIIc of Hassler's nomenclature, corresponding to Brodmann's layer IVb) were predominantly pyramidal, although many spiny multipolar (stellate) cells were also found. The average basal dendritic field area of pyramidal neurons in sublamina IIIc of V1 was significantly smaller than that in the homologous layer of V2, within the cytochrome oxidase-rich thick stripes. Furthermore, the average basal dendritic field areas of V1 and V2 pyramidal neurons were significantly smaller than those of neurons in MT, LIPv and area 7a. There was no difference in basal dendritic field area between layer III pyramidal neurons in areas MT, LIPv and 7a. While the shape of most basal dendritic fields was circularly symmetrical in the dimension tangential to the cortical layers, there were significant biases in complexity, with dendritic branches tending to cluster along particular axes. Sholl analysis revealed that the dendritic fields of neurons in areas MT, LIPv and 7a were significantly more complex (i.e. had a larger number of branches) than those of V1 or V2 neurons. Analysis of basal dendritic spine densities revealed regional variations along the dendrites, with peak densities being observed 40-130 microns from the cell body, depending on the visual area. The peak spine density of layer III pyramidal neurons in V1 was lower than that observed in V2, MT or LIPv, which were all similar. Pyramidal neurons in area 7a had the greatest peak spine density, which was on average 1.7 times that found in V1. Calculations based on the average spine density and number of dendritic branches at different distances from the cell body demonstrated a serial increase in the total number of basal dendritic spines per neuron at successive stations of the occipitoparietal pathway. Our observations, comparing dendritic fields of neurons in the homologous cortical layer at different levels of a physiologically defined 'stream', indicate changes in pyramidal cell morphology between functionally related areas. The relatively large, complex, spine-dense dendritic fields of layer III pyramidal cells in rostral areas of the occipitoparietal pathway allow these cells to sample a greater number of more diverse inputs in comparison with cells in 'lower' areas of the proposed hierarchy.      

Cell type- and region-specific expression of neurogranin mRNA in the cerebral cortex of the macaque monkey

Neurogranin is a postsynaptic substrate for protein kinase C (PKC). It has been identified in the central nervous system, and the expression has been related to postsynaptic plasticity. Using non-radioactive in situ hybridization histochemistry, we investigated whether mRNA expression of neurogranin varied among the cerebral region and cell types. In most areas of the neocortex excluding area OC (the primary visual area), intense signals were observed in the pyramidal cells in layers III, V and VI. In area OC, intense signals were observed in layers IV as well as layers III and VI. We previously showed that intense signals for GAP-43, a presynaptic PKC substrate, were observed in relay neurons of the lateral geniculate nucleus. From this result and the present result in area OC, we conclude that both pre- and postsynaptic PKC substrates (GAP-43 and neurogranin) are abundant in the geniculocortical synapses. In the hippocampus, intense signals were observed in the pyramidal cells in the subiculum. Taken together with our previous study showing intense signals for GAP-43 in Ammon's horn, the result indicates that both PKC substrates are abundant in the connections between neurons in Ammon's horn and in the subiculum.     

Functional diversity of layer IV spiny neurons in rat somatosensory cortex: quantitative morphology of electrophysiologically characterized and biocytin labeled cells

Previous analyses of the spiny layer IV neurons have almost exclusively focused on spiny stellate cells. Here we provide detailed morphological data characterizing three subpopulations of spiny neurons in slices of adolescent rats: (i) spiny stellate cells (58%), (ii) star pyramidal cells (25%) and (iii) pyramidal cells (17%), which can be distinguished objectively by the preferential orientation of their dendritic stems. Spiny stellate cells lacked an apical dendrite and frequently confined their dendritic and axonal arbors to the respective column. Star pyramidal and pyramidal cells possessed an apical dendrite, which reached the supragranular layers. Their axonal arbors were similar, showing both a columnar component and transcolumnar branches with direct transbarrel projections. However, a small fraction of star pyramidal cells possessed few or even no transcolumnar branches. Electrophysiologically, all three types of neurons were either regular-spiking or intrinsically burst-spiking without a significant relation to the morphological subtypes. The basic synaptic properties of thalamic inputs were also independent of the type of target layer IV spiny neuron. All remained subthreshold and showed paired-pulse depression. In conclusion, the columnar axonal arborization of spiny stellate cells is supplemented by a significant oblique to horizontal projection pattern in pyramidal-like neurons. This offers a structural basis for either segregation or early context-dependent integration of tactile information, in a cell-type specific manner.     

Relationships of local inhibitory and excitatory circuits to orientation preference maps in ferret visual cortex.

The contribution and precise role of intracortical circuits in generating orientation tuned responses in visual cortical neurons is still controversial. To address this question, the relationship between excitatory and inhibitory synaptic connections and orientation maps in ferret striate cortex was investigated by combining in vivo optical imaging and in vitro scanning laser photostimulation. Excitatory and inhibitory inputs to pyramidal cells originated preferentially from regions with similar orientation preference. Prominent cross-orientation inhibition was not observed, arguing against cross-orientation models of orientation selectivity. The tuning of inhibitory inputs was significantly broader in both layer 2/3 and layer 5/6 pyramidal neurons compared to the tuning of excitatory inputs. Local excitatory inputs were more prominent in the 0-20 degrees tuning difference range between pre- and postsynaptic cells than inhibitory inputs, whereas inhibition dominated in the 20-40 degrees tuning difference range. These differences in tuning of excitatory and inhibitory inputs onto individual cells are consistent with the predictions of recurrent models of orientation selectivity.     

A neural model of how horizontal and interlaminar connections of visual cortex develop into adult circuits that carry out perceptual grouping and learning

A neural model suggests how horizontal and interlaminar connections in visual cortical areas V1 and V2 develop within a laminar cortical architecture and give rise to adult visual percepts. The model suggests how mechanisms that control cortical development in the infant lead to properties of adult cortical anatomy, neurophysiology and visual perception. The model clarifies how excitatory and inhibitory connections can develop stably by maintaining a balance between excitation and inhibition. The growth of long-range excitatory horizontal connections between layer 2/3 pyramidal cells is balanced against that of short-range disynaptic interneuronal connections. The growth of excitatory on-center connections from layer 6-to-4 is balanced against that of inhibitory interneuronal off-surround connections. These balanced connections interact via intracortical and intercortical feedback to realize properties of perceptual grouping, attention and perceptual learning in the adult, and help to explain the observed variability in the number and temporal distribution of spikes emitted by cortical neurons. The model replicates cortical point spread functions and psychophysical data on the strength of real and illusory contours. The on-center, off-surround layer 6-to-4 circuit enables top-down attentional signals from area V2 to modulate, or attentionally prime, layer 4 cells in area V1 without fully activating them. This modulatory circuit also enables adult perceptual learning within cortical area V1 and V2 to proceed in a stable way.     

Integration of local inputs in visual cortex

In mammalian visual cortex, local connections are ubiquitous, extensively linking adjacent neurons of all types. In this study, optical maps of intrinsic signals and responses from single neurons were obtained from the same region of cat visual cortex while the effectiveness of the local cortical circuitry was altered by focally disinhibiting neurons within a column of known orientation preference. Maps of intrinsic signals indicated that local connections provide strong and functional subthreshold inputs to neighboring columns of other orientation preferences, altering the observed orientation preference to that of the disinhibited column. However, measuring the suprathreshold response using single-cell recordings revealed only mild changes of preferred orientation over the affected region. Because strongly tuned subthreshold inputs from cortex only marginally affect the tuning of a cortical cell's output, it is concluded that local cortical inputs are integrated weakly compared to geniculate inputs. Such circuitry potentially allows for the normalization of responses across a wide range of input activity through local averaging.      

Morphological variation of layer III pyramidal neurones in the occipitotemporal pathway of the macaque monkey visual cortex

We compared the morphological characteristics of layer III pyramidal neurones in different visual areas of the occipitotemporal cortical 'stream', which processes information related to object recognition in the visual field (including shape, colour and texture). Pyramidal cells were intracellularly injected with Lucifer Yellow in cortical slices cut tangential to the cortical layers, allowing quantitative comparisons of dendritic field morphology, spine density and cell body size between the blobs and interblobs of the primary visual area (V1), the interstripe compartments of the second visual area (V2), the fourth visual area (V4) and cytoarchitectonic area TEO. We found that the tangential dimension of basal dendritic fields of layer III pyramidal neurones increases from caudal to rostral visual areas in the occipitotemporal pathway, such that TEO cells have, on average, dendritic fields spanning an area 5-6 times larger than V1 cells. In addition, the data indicate that V1 cells located within blobs have significantly larger dendritic fields than those of interblob cells. Sholl analysis of dendritic fields demonstrated that pyramidal cells in V4 and TEO are more complex (i.e. exhibit a larger number of branches at comparable distances from the cell body) than cells in V1 or V2. Moreover, this analysis demonstrated that the dendrites of many cells in V1 cluster along specific axes, while this tendency is less marked in extrastriate areas. Most notably, there is a relatively large proportion of neurones with 'morphologically orientation-biased' dendritic fields (i.e. branches tend to cluster along two diametrically opposed directions from the cell body) in the interblobs in V1, as compared with the blobs in V1 and extrastriate areas. Finally, counts of dendritic spines along the length of basal dendrites revealed similar peak spine densities in the blobs and the interblobs of V1 and in the V2 interstripes, but markedly higher spine densities in V4 and TEO. Estimates of the number of dendritic spines on the basal dendritic fields of layer III pyramidal cells indicate that cells in V2 have on average twice as many spines as V1 cells, that V4 cells have 3.8 times as many spines as V1 cells, and that TEO cells have 7.5 times as many spines as V1 cells. These findings suggest the possibility that the complex response properties of neurones in rostral stations in the occipitotemporal pathway may, in part, be attributed to their larger and more complex basal dendritic fields, and to the increase in both number and density of spines on their basal dendrites.      

Numerical Relationships between Geniculocortical Afferents and Pyramidal Cell Modules in Cat Primary Visual Cortex

An analysis has been made of the quantitative data availableon the number of pyramidal cell modules of layer IV neurons,and of geniculocortical axons and their synapses in cat striatecortex. It is found that the convergence of geniculocorticalafferents upon any one pyramidal cell module is enormous, sincein any one location there is overlap between 360–540 X-axonsand 300–540 Y-axons. In total, the X- and Y-axonal arborsprovide some 1640 x 10⁶ synapses to area 17, which is equivalentto a ratio of 160–200 synapses per layer IV neuron. Thesevalues assume that geniculocortical terminals synapse only withthe spiny stellate cells of layer IV. The values are reducedto 100–125 per spiny stellate cell when account is takenof the synapses that involve the dendrites that enter layerIV from neurons with cell bodies in other layers. Since eachlayer IV neuron receives some 2500 asymmetric synapses, thismeans that only 5% of the total excitatory input to a layerIV neuron seems to be provided by the geniculocortical afferents.Further, if the boutons in the geniculocortical axonal arborsare distributed homogeneously across layer IV, each axon couldonly provide one synapse to about one in four of the layer IVneurons encompassed by its plexus. It may be, however, thatinstead of being spread evenly, boutons in individual arborsconverge upon individual neurons to supply a number of synapsesto them. But even so, it seems unlikely that any individualgeniculate axon could dominate the activity of a particularcortical neuron.     

Dynamic properties of excitatory synaptic connections involving layer 4 pyramidal cells in adult rat and cat neocortex

To investigate the properties of excitatory connections between layer 4 pyramidal cells and whether these differed between rat and cat, paired intracellular recordings were made with biocytin filling in slices of adult neocortex. These connections were also compared with those from layer 4 spiny cells to layer 3 pyramids and connections between layer 3 pyramids. Connectivity ratios for layer 4 pyramid-pyramid pairs (1:14 cat, 1:18 rat) appeared lower than for the other types of connections studied in parallel, but excitatory postsynaptic potential (EPSP) amplitudes and time course were not significantly different either between species or across types of connection. Layer 4 pyramids targeted postsynaptic basal dendrites in both species, whether the pyramidal target was in layer 4 or layer 3. Within layer 4, relationships between mean EPSP amplitude, numbers of putative contacts, and distance between connected pairs indicated a rapid decline in connectivity strength with distance, equivalent to 3.4 mV and 10 synapses per 100 microm separation, from a maximum of 4 mV and 10 synapses at 0 microm. However, a subset, of burst-firing layer 4 pyramids, appeared to make no connections with other layer 4 spiny cells. Second EPSPs were depressed by 36% in rat and 28% in cat relative to first EPSPs at interspike intervals <15 ms. Subsequent EPSPs in brief trains were further depressed. Depression was predominantly presynaptic in origin. Recovery from depression could not be described adequately by a simple exponential for individual connections; it included peaks and troughs with periodicities of 10-15 ms. Complex relationships between the first 2 interspike intervals and third EPSP amplitude were also apparent in all connections so studied. Large third EPSPs followed specific combinations of first and second interspike intervals so that increasing, or decreasing, one without changing the other resulted in a smaller third EPSP. Finally, the outputs of layer 4 spiny cells to layer 3 exhibited partial recovery from depression during longer high-frequency trains, a property not apparent in the other connections studied.     

Prenatal development of calbindin D-28K in human visual cortex

The distribution of the calcium-binding protein calbindin D-28K (CB) was investigated in human fetal primary visual cortex. CB is present in Cajal-Retzius cells of layer I, in sparse neurons of the ventricular and intermediate zones (VZ, IZ), and in tangential fibres in IZ by 15 weeks (W) of gestation. Cajal-Retzius cells lose their staining by 30W. CB appears in layers II-VI mainly from 26W, following an inside-outside sequence. Until 34W, CB labelling is in somata and neuropil located primarily in layers IVA, IVC and V. Then reactive perikarya and puncta increase in layers II-IVA and deep IVB and C, but are reduced in infragranular layers from 34W to term. From 30W positive somata form clusters in the cell-rich bands in layers IV and V and labelled neuropil in layers III and IV has a periodic pattern from 34W. Also from 34W, numerous lightly reactive pyramidal cells are present in layers II to IVA in primary, but not secondary, visual cortex. Our results show precocious expression of CB before full laminar differentiation of the cortex and that some of this expression is transient.      

Confocal Microscopic Study of the Dendritic Organization of Patchy, Intrinsic Neurons in Area 18 of the Cat

The intrinsic, horizontally projecting neurons in visual cortexare organized into discrete clusters, or patches. While earlierstudies focused on the organization of the patches and theirrelationship to functional properties, such as orientation tuningand binocularity, little is known about the detailed morphologyof the neurons in these patches. We retrogradely labeled patchesof local, intracortical neurons (local patch neurons) by invivo injections of fluorescent dextrans into area 18, then iontophoresed Lucifer yellow into prelabeled cells in lightly fixedcortical slices, and examined the dendritic morphology of localpatch neurons in layer 2/3 of area 18 by confocal laser scanningmicroscopy. Most of the neurons we examined were spiny pyramidalneurons, including modified pyramids, small to moderately largestandard pyramids, and star pyramids. Smooth, multipolar cellsof the basket and bipolar cell types were also present. Thebasal dendritic trees of more than 60% of the local patch pyramidalneurons in layer 2/3 displayed mediolaterally elongated dendriticfields. This finding appears to be specific for pyramidal neurons,as our sample of smooth, multipolar neurons did not show thistrend.     

Different inhibitory synaptic input patterns in excitatory and inhibitory layer 4 neurons of ferret visual cortex

The synaptic mechanisms underlying the generation of orientation and direction selectivity in layer 4 of the primary visual cortex are still largely unclear. Previous in vivo work has shown that intra-cortical inhibition plays a major role in generating the properties of orientation and direction selectivity. Excitatory and inhibitory cortical neurons differ in their receptive field properties: excitatory neurons tend to be orientation- and direction-selective, inhibitory neurons tend to be orientation-, but not direction-selective. Here we have compared the relationship between direction preference maps recorded in vivo and synaptic input maps recorded in vitro from excitatory and inhibitory stellate cells in layer 4 of ferret visual cortex. Our goal was to test whether the differences in direction tuning between these cell populations might result from different inhibitory connectivity patterns. We found that excitatory neurons, which are direction tuned in vivo, receive approximately 50% of their inhibitory inputs from cortical regions of opposite direction preference whereas inhibitory cells, which are not or poorly direction tuned, receive only very few inputs from regions of opposite direction preference. This confirms that inhibitory connections arising in cortical regions of opposite direction preference may be required to create or strengthen direction tuning in their target neurons. Thus, differences in intracortical inhibitory circuit patterns may underlie the differences in receptive field properties observed between excitatory and inhibitory neurons in vivo.     

A study of pyramidal cell structure in the cingulate cortex of the macaque monkey with comparative notes on inferotemporal and primary visual cortex

Recent studies have revealed a marked degree of variation in the pyramidal cell phenotype in visual, somatosensory, motor and prefrontal cortical areas in the brain of different primates, which are believed to subserve specialized cortical function. In the present study we carried out comparisons of dendritic structure of layer III pyramidal cells in the anterior and posterior cingulate cortex and compared their structure with those sampled from inferotemporal cortex (IT) and the primary visual area (V1) in macaque monkeys. Cells were injected with Lucifer Yellow in flat-mounted cortical slices, and processed for a light-stable DAB reaction product. Size, branching pattern, and spine density of basal dendritic arbors was determined, and somal areas measured. We found that pyramidal cells in anterior cingulate cortex were more branched and more spinous than those in posterior cingulate cortex, and cells in both anterior and posterior cingulate were considerably larger, more branched, and more spinous than those in area V1. These data show that pyramidal cell structure differs between posterior dysgranular and anterior granular cingulate cortex, and that pyramidal neurons in cingulate cortex have different structure to those in many other cortical areas. These results provide further evidence for a parallel between structural and functional specialization in cortex.     

Morphometric analysis of the columnar innervation domain of neurons connecting layer 4 and layer 2/3 of juvenile rat barrel cortex

We have investigated the dendritic and axonal morphology of connected pairs of L4 spiny neurons and L2/3 pyramidal cells in rat barrel cortex. The 'projection' field of the axons of L4 spiny neurons in layers 2/3, 4 and 5 has a width of 400-500 microm thereby defining an anatomical barrel-column. In layer 2/3, the averaged axonal 'projection' field of L4 spiny neurons together with the dendritic 'receptive' field of the connected L2/3 pyramidal cells form a mostly column-restricted anatomical L4-to-L2/3 'innervation domain' that extends 300-400 microm and includes mostly basal dendrites. In the L4-to-L2/3 innervation domain a single L4 spiny neuron contacts approximately 300-400 pyramidal cells while in the L4-to-L4 innervation domain it contacts approximately 200 other L4 spiny neurons. Similarly approximately 300-400 L4 spiny neurons converge onto a single pyramidal cell and approximately 200 L4 spiny neurons innervate another L4 spiny neuron. The L2/3 pyramidal cell axon has a vertical projection field spanning all cortical layers, and a long-range horizontal field in layers 2/3 (width 1,100-1,200 microm) and 5 (700-800 microm) projecting across column borders. The results suggest that the flow of excitation within a barrel-column is determined by the largely columnar confinement of the L4-to-L4 and L4-to-L2/3 innervation domains. A whisker deflection activates approximately 140 L4 spiny neurons that will generate EPSPs in most barrel-related L2/3 pyramidal cells of a principal whisker column. The translaminar synaptic transmission to layer 2/3 and the axonal projection fields of L2/3 pyramidal cells are the major determinants of the dynamic, multi-columnar map in which a single whisker deflection is represented in the cortex.     

Laminar specificity of intrinsic connections in Broca's area

Broca's area and its right hemisphere homologue comprise 2 cytoarchitectonic subdivisions, FDgamma and FCBm of von Economo C and Koskinas GN (1925, Die Cytoarchitektonik der Hirnrinde des erwachsenen Menschen. Vienna/Berlin [Germany]: Springer). We report here on intrinsic connections within these areas, as revealed with biotinylated dextran amine and 1,1'-dioctadecyl-3,3,3',3'-tetramethylindocarbocyanine perchlorate tracing in postmortem human brains. Injections limited to supragranular layers revealed a complex intrinsic network of horizontal connections within layers II and III spreading over several millimeters and to a lesser extent within layers IV, V, and VI. Ninety percent of the retrogradely labeled neurons (n = 734) were in supragranular layers, 4% in layer IV, and 6% in infragranular layers; most were pyramids and tended to be grouped into clusters of approximately 500 microm in diameter. Injections involving layer IV revealed extended horizontal connections within layers I-IV (up to 3.7 mm) and to a lesser extent in layers V and VI. Injections limited to the infragranular layers revealed horizontal connections mainly limited to these layers. Thus, intrinsic connections within Broca's area display a strong laminar specificity. This pattern is very similar in areas FDgamma and FCBm and in the 2 hemispheres.     

Laminar patterns of local excitatory input to layer 5 neurons in macaque primary visual cortex

Layer 5 neurons in primary visual cortex make putative reciprocal feedback connections to the superficial layers. To test this hypothesis, we employed scanning laser photostimulation combined with intracellular dye injection to examine local functional excitatory inputs to and axonal projections from individual layer 5 neurons in brain slices from monkey V1. In contrast with previous studies of other V1 neurons, layer 5 neurons received significant input from nearly all of the cortical layers, suggesting individual layer 5 cells integrate information from a broad range of input sources. Nevertheless relative strengths of laminar inputs varied across neurons. Cluster analysis of relative strength of laminar inputs to individual layer 5 neurons revealed four discrete clusters representing recurring input patterns; each cluster included both excitatory and inhibitory neurons. Twenty-five of 40 layer 5 neurons fell into two clusters, both characterized by very strong input from superficial layers. These input patterns are consistent with layer 5 neurons providing feedback to superficial layers. The remaining 15 neurons received stronger input from deep layers. Differences in input from layer 4Calpha versus 4Cbeta also suggest specific associations of the magnocellular and parvocellular visual pathways, with populations receiving stronger input from deep versus superficial cortical layers.