Anatomical banding of intrinsic connections in striate cortex of tree shrews (Tupaia glis)

The intrinsic connectivity of striate cortex was investigated by injecting horseradish peroxidase (HRP) into this area in tree shrews. Such HRP injections demonstrated periodically organized, stripelike connections within area 17. These stripes occur in layers I–IIIA and consist of a small number of retrogradely filled neurons, some clearly pyramidal, together with HRP-labeled axon terminals. HRP-filled axon trunks run between labeled stripes, interconnecting adjacent and distant regions of the stripe pattern. Correlation with Golgi-stained tissue suggests that these stripes are horizontally interconnected by pyramidal neurons with long intracortical axon collaterals (followed for distances over 1 mm form the soma). The HRP-labeled stripes measure about 230 μm in width, with a center-to-center repeat distance of 450–500 μm. They have been mapped over an 8 mm² area of striate cortex and would thus seem capable of effecting lateral interactions over considerable portions of the retinotopic map. In their dimensions and overall pattern, these anatomical stripes resemble the 2-deoxyglucose (2-DG) bands resulting from visual stimulation of tree shrews with stripes of a single orientation. While the functional role of the HRP-labeled stripes is unclear, their similarities with the 2-DG pattern raise the intriguing possibility that they may be related to orientation selectivity. The striking regularity of these extensive lateral interconnections emphasizes the importance of horizontal intralaminar connections within the cortex.     

A reticular pattern of intrinsic connections in primate area V2 (area 18)

A system of periodic intrinsic connections is demonstrated in area V2 (area 18) of squirrel and macaque monkeys by large injections of tritiated amino acids, horseradish peroxidase (HRP), and fluorescent latex beads. These connections originate from pyramidal neurons concentrated in layers 3 and 5. Terminations occur in all cortical layers, largely coextensive with labeled neurons but more restricted in layer 4. This multilaminar distribution contrasts with the mainly supragranular localization of periodic intrinsic connections in V1 (area 17), and may imply a close interaction, in V2, of periodic intrinsic connections with pulvinocortical, as well as with corticocortical terminations (concentrated, respectively, in layers 3 and 5, and in lower 3 and 4). As in V1, the tangential configuration of these connections in V2 is reticular or latticelike, and is detectable for 2.5-3.0 mm from an injection site of HRP, 3H amino acids, or latex beads. Cross-sectional widths of labeled regions vary from 250 to 800 micron in squirrel monkey and from 400 to 1,000 micron in macaque, depending on which portion of the lattice is measured. When periodic intrinsic connections are compared with stripes labeled histochemically by cytochrome oxidase (CO), no clear relationship is obvious between the two systems. This result contrasts with the orderly tangential alignment reported between CO-reactive zones in V2 and certain extrinsic connections; namely, pulvinocortical terminations (Livingstone and Hubel, '82) and clusters of neurons projecting to area V4 (DeYoe and Van Essen, '84). Other extrinsic connections, however, such as backgoing connections from V2 to V1, do not seem to have a periodic distribution. Thus, although some discontinuous cortical connections relate to each other in a precise mosaic fashion, intrinsic and some extrinsic connections may observe different modes of organization.     

Cortical connections of areas 17 (V-I) and 18 (V-II) of squirrels

Connections of visual cortex in squirrels were investigated by placing WGA-HRP injections, and in some cases fluorescent dyes, into area 17 (V-I) or area 18 (V-II). Results were related to architectonic fields determined in brain sections cut parallel to the surface of manually flattened cortex and to limited microelectrode mapping data. Injections in area 17 provided evidence for 1) a patchy pattern of horizontal intrinsic connections extending 1-2 mm from the injection site; 2) uneven, widely distributed connections with area 18 (V-II) and adjoining occipital-temporal (OT) cortex; and 3) callosal connections of large portions of area 17 with the 17/18 border zone. While restricted locations in area 17 had uneven interconnections over several mm of area 18, more rostral locations in area 17 related to more rostral locations in area 18, demonstrating a topographic tendency. Injections in area 18 revealed 1) zones of discontinuous connections with area 17 that followed a topographic pattern, 2) patches of intrinsic connections that spread over distances of up to 6-8 mm from the injection site; 3) two zones of uneven connections with OT cortex suggesting the locations of at least two visual areas, OTr and OTc; 4) connections with limbic cortex rostromedial to areas 17 and 18; 5) sparse connections with regions of temporal cortex lateral to OT; and 6) uneven callosal connections with area 18 and OT cortex. The widespread and unevenly distributed intrinsic callosal interconnection patterns of areas 17 and 18 contrast with the restricted excitatory receptive fields of neurons and the retinotopic patterns of representation in these fields. Although physiological evidence is presently lacking, the patchy connections suggest that areas 17 and 18 in squirrels are modularly organized.     

Interhemispheric connections in the visual cortex of the squirrel monkey (Saimiri sciureus)

The callosal connections within the posterior parietal and occipital cortices were studied in the squirrel monkey with horseradish peroxidase tracing techniques. The data were evaluated with particular emphasis on the relationship of major callosal connections along the 17-18 border. The overall pattern of callosal connections in the squirrel monkey also was compared with callosal patterns in other New World simians. Our results show that the dense band of callosal connections along the 17-18 border in the squirrel monkey differs from the connections observed in other New World monkeys in that it is virtually confined to area 18 and avoids area 17. In addition to a continuous band of callosal connections in area 18 that parallels the 17-18 border, rostral extensions of the band are oriented perpendicular to the 17-18 border and present an obvious periodicity. The remaining parieto-occipital cortex contains a complex pattern of callosal connections that is strikingly similar to patterns reported for other New World monkeys. Thus, it is likely that the dorsolateral extrastriate visual cortex in the squirrel monkey is organized in a manner similar to that found within other New World monkeys.     

Columnar specificity of intrinsic horizontal and corticocortical connections in cat visual cortex

A prominent and stereotypical feature of cortical circuitry in the striate cortex is a plexus of long-range horizontal connections, running for 6-8 mm parallel to the cortical surface, which has a clustered distribution. This is seen for both intrinsic cortical connections within a particular cortical area and the convergent and divergent connections running between area 17 and other cortical areas. To determine if these connections are related to the columnar functional architecture of cortex, we combined labeling of the horizontal connections by retrograde transport of rhodamine-filled latex microspheres (beads) and labeling of the orientation columns by 2-deoxyglucose autoradiography. We first mapped the distribution of orientation columns in a small region of area 17 or 18, then made a small injection of beads into the center of an orientation column of defined specificity, and after allowing for retrograde transport, labeled vertical orientation columns with the 2-deoxyglucose technique. The retrogradely labeled cells were confined to regions of orientation specificity similar to that of the injection site, indicating that the horizontal connections run between columns of similar orientation specificity. This relationship was demonstrated for both the intrinsic horizontal and corticocortical connections. The extent of the horizontal connections, which allows single cells to integrate information over larger parts of the visual field than that covered by their receptive fields, and the functional specificity of the connections, suggests possible roles for these connections in visual processing.     

Cortical connections of area 17 in tree shrews

In order to better understand the organization of extrastriate cortex in tree shrews, injections in area 17 of wheat germ agglutinin or tritiated proline were used to reveal an intrinsic pattern of connections, ipsilateral connections with area 18 and two other subdivisions of cortex, and callosal connections with areas 17 and 18 of the opposite cerebral hemisphere. Areal patterns of connections were best seen in sections cut parallel to the surface of flattened cortex. Within area 17, periodic foci of labeled terminations and cells extended from and surrounded injection sites as described by Rockland et al. ('82). Single injections produced multiple foci of labeled terminations and cells in area 18. The foci tended to fuse into short bands that sometimes crossed the width of area 18. Double injections produced more foci, and multiple injections tended to produce more continuous regions of label. An overall retinotopic pattern was evident with rostral area 17 connected to rostral area 18 and caudal area 17 connected to caudal area 18. Terminations extended through layers II-VI, with some increase in density in layer IV. Cells in area 18 projecting back to area 17 were in layers III and V. The injections also allowed identification of previously undefined subdivisions of visual cortex in temporal cortex immediately adjoining area 18. Dense reciprocal connections were observed in a 13 mm2 oval of cortex on the lateral border of the middle section of area 18 that we define as the temporal dorsal area, TD. Connections indicate a crude topographic organization with lower field represented rostrally and upper field caudally. Inputs were most dense in the middle cortical layers, and labeled cells were supragranular, and less frequently, infragranular. A 10-mm2 oval of cortex near the posterior edge of the hemisphere, the temporal posterior area (TP), contained labeled cells after area 17 injections, but terminal labeling was only obvious in the dorsal part. Single injections sometimes produced quite separate dorsal and ventral zones of label in TP, suggesting a small separate dorsal division. A crude retinotopic order appears to exist within ventral TP, with the lower field most ventral. Labeled cells were largely supragranular. A fourth zone of ipsilateral connections was in posterior limbic cortex bordering area 17 on the ventromedial surface of the cerebral hemisphere. The callosal connections were reciprocal and included regions 1 mm wide on either side of the area 17 and area 18 border. Callosal connections were rougly homotopic. Callosal terminations included superficial layers, and projecting cells were both supragranular and infragranular.(ABSTRACT TRUNCATED AT 400 WORDS)     

Organization of local axon collaterals of efferent projection neurons in rat visual cortex

We have studied the laminar origins of local long-range connections within rat primary visual cortex (area 17), by using retrograde tracing of nerve cell bodies with fluorescent markers. Injections throughout the thickness of cortex produce distinct laminar labeling patterns which indicate that a substantial number of cells in layers 2/3, 5, and 6 have wide local axon collateral arbors, while the local arbors of layer 4 cells are much narrower. Double labeling experiments which combined area 17 injections with injections into different projection targets of area 17 (opposite area 17, area 18a, and area 18b) show that many cortico-cortically projecting cells make widespread projections within area 17. In contrast, the overwhelming majority of subcortically projecting cells have narrow collateral arbors within area 17. Anterograde tracing of local projections within areas 17 with the lectin Phaseolus vulgaris leucoagglutinin shows an extensive system of horizontally running fibers which terminate in distinct 0.15-0.25 mm wide clusters up to 1.8 mm from the injection site. On horizontal sections the termination pattern resembles a closely spaced lattice. The results indicate that cortico-cortically projecting cells provide for long-range interactions between distant points of the visuotopic map, while subcortically projecting cells mediate information within a cortical column. Interestingly, subcortically projecting cells differ functionally from cortico-cortically projecting cells in that they are not orientation selective (Klein et al., Neurosci. 17:57-78, '86; Mangini and Pearlman, J. Comp. Neurol. 193:203-222, '80; Simmons and Pearlman, J. Neurophysiol. 50:838-848, '83). We therefore suggest that cortico-cortically projecting cells with wide collateral arbors are orientation selective and that clustered long-range projections within area 17 connect columns with similar functional specificity.     

Anatomical consequences of long-term monocular eyelid closure on lateral geniculate nucleus and striate cortex in squirrel monkey

The effects of long-term monocular deprivation on the geniculostriate system in squirrel monkeys were studied with neuroanatomical methods. Four neonates were visually deprived by monocular eyelid suture during their first 10 days of life and survived from 9 to 40 months. In the lateral geniculate nucleus (LGN), deprivation resulted in severe cell size changes. Neurons in the deprived laminae were smaller compared to those in the undeprived laminae. Deprivation left the reciprocal connections between LGN and striate cortex intact: After horseradish peroxidase (HRP) injections into striate cortex, retrogradely transported enzyme labeled a wedge of neurons in deprived and undeprived LGN laminae; anterogradely transported HRP filled preterminal and terminal axons in this wedge. Following 3H-proline injections into the deprived eye for transneuronal transport, autoradiography showed in the ipsilateral striate cortex a silver grain distribution over most of layer IVc similar to that in normal squirrel monkeys, except for a small strip in the anterior calcarine fissure. Here, a few, irregularly spaced "patches" of higher grain density occurred deep in layer IVc. Layer IVc of contralateral area 17 was also uniformly labeled over most of its extent, except for a very few and inconspicuous accumulations of slightly increased silver grains. After visual stimulation of the deprived eye, the 14C-2-deoxyglucose method showed in the contralateral striate cortex some alternating "patches" of higher uptake superimposed on the heavy labeling in layer IVc. Layer IVc in the ipsilateral cortex was more uniformly labeled. Regularly spaced arrays of labeled "puffs" in layers II/III were present in both hemispheres. Cytochrome oxidase staining showed no change in the distribution pattern of the enzyme in the deprived monkeys from the basic pattern of normal adults. No changes in cell sizes were found in layer IVc in cresyl-violet-acetate-stained sections. These results lead to the conclusion that in area 17 of squirrel monkeys there is no distinct segregation of inputs from the two eyes into anatomically discrete ocular dominance columns and they support the view of a predominantly binocular organization of area 17.     

Feedback connections to ferret striate cortex: direct evidence for visuotopic convergence of feedback inputs

Interareal feedback connections are a fundamental aspect of cortical architecture, yet many aspects of their organization and functional relevance remain poorly understood. Previous studies have investigated the topography of feedback projections from extrastriate cortex to macaque area 17. We have extended this analysis to the ferret. We made restricted injections of cholera toxin B (CTb) into ferret area 17 and mapped the distribution of retrogradely labeled cells in extrastriate cortex. In addition to extensive label spreading within area 17, we found dense cell label in areas 18, 19, and 21 and the suprasylvian cortex and sparser connections from the lateral temporal and posterior parietal cortex. We made extensive physiological assessments of magnification factors in the extrastriate visual cortex and used these measures to convert the spread of labeled cortex in millimeters into a span in degrees of visual field. We also directly measured the visuotopic extents of receptive fields in the regions containing labeled cells in cases in which we made both CTb injections and physiological recordings in the same animals; we then compared the aggregate receptive field (ARF) of the labeled region in each extrastriate area with that of the injection site. In areas 18, 19, and 21, receptive fields of cells in regions containing labeled neurons overlapped those at the injection site but spanned a greater distance in visual space than the ARF of the injection site. The broad visuotopic extent of feedback connections is consistent with the suggestion that they contribute to response modulation by stimuli beyond the classical receptive field.     

Interhemispheric connections of visual cortex of owl monkeys (Aotus trivirgatus), marmosets (Callithrix jacchus), and galagos (Galago crassicaudatus)

Interhemispheric connections of visual cortex were studied in owl monkeys, marmosets, and galagos after multiple injections of horseradish peroxidase into one cerebral hemisphere. Areal patterns of connections were revealed in sections of cortex that was flattened and cut parallel to the surface. Results were related to the locations of known visual areas, especially in owl monkeys, in which more visual areas have been established. The connection patterns in owl monkeys and marmosets are very similar, suggesting that the organization of visual cortex differs little in these two New World simians. Galagos have a basically similar pattern, but the connections are more widespread. In all three primates, connections are not restricted to cortex representing the line of decussation of the retina, and even striate cortex has connections displaced from the border. These connections extend up to 2 mm into area 17 in owl monkeys, and they are most extensive in galagos, where they form foci that are coextensive with regions of high cytochrome oxidase activity. Connections are concentrated in the caudal half of area 18, but protrusions of connections cross of the width of the field. The middle temporal visual area (MT) has unevenly distributed connections throughout, with some increase in density along the border. The dorsomedial visual area (DM) of owl monkeys has connections restricted to the rostral border, and a similar region of sparse connections identifies the probable location of DM in marmosets and galagos. Caudal parts of the dorsolateral visual area (DL) of owl monkeys have dense interhemispheric connections. Other visual areas are characterized by unevenly distributed clumps of connections, suggesting that functions are not uniformly distributed, and that semiregular processing modules exist. The results indicate that most extrastriate visual neurons are subject to interhemispheric influences and support the conclusion that callosal connections are functionally heterogeneous.     

Topography of developing thalamic and cortical pathways in the visual system of the cat

Adult patterns of connectivity could emerge during development by a process of selective elimination from an earlier, more widespread, connectivity. We have addressed this issue by examining the topography of developing projections to area 17 in the cat. At different postnatal ages, paired injections of the retrograde tracers diamidino yellow and fast blue were made in area 17. Interinjection separations were carefully controlled and the spatial distribution of the two populations of labelled neurones investigated. Projections to the striate cortex from the lateral geniculate nucleus, area 18, as well as connections intrinsic to area 17 were analysed quantitatively with a graphic method that uses a two-dimensional model of the projection. This allows two parameters of the projection to be calculated: the divergence (the spatial extent of area 17 contacted by an infinitely small region of an afferent structure) and the convergence (the extent of an afferent structure that projects to an infinitely small region of area 17). During postnatal development, the bulk of the connections making up the geniculostriate and corticocortical pathways showed no variation either in their convergence and divergence. However, the projection of area 18 to area 17 and the intrinsic area 17 connections (but not the geniculostriate projection) in the 3–15-day-old kittens were each found to contain a small subpopulation of widely scattered neurones with widespread axonal trajectories. These results, showing that many initially formed connections display a high degree of topographical order, are discussed in terms of the control mechanisms specifying axonal trajectories during development. © 1994 Wiley-Liss, Inc.     

Effects of neonatal enucleation on the organization of callosal linkages in striate cortex of the rat

Lewis and Olavarria ([1995] J. Comp. Neurol. 361:119–137) showed that the mediolateral organization of callosal linkages differs markedly between medial and lateral regions of striate cortex in the rat. Thus, callosal fibers originating from medial regions of striate cortex interconnect loci that are mirror-symmetric with respect to the midsagittal plane. In contrast, fibers from lateral regions of striate cortex show a reversed pattern of connections: tracer injections into the 17/18a border produce retrograde cell labeling in regions medial to the contralateral 17/18a border, whereas injections placed somewhat medial to the 17/18a border label cells located at the contralateral 17/18a border. Based on the interpretation that callosal fibers from lateral striate cortex connect retinotopically corresponding loci (Lewis and Olavarria [1995] J. Comp. Neurol. 361:119–137) we propose here that the development of the reversed pattern of connections in lateral portions of striate cortex is guided by activity-dependert cues originating from spontaneously active ganglion cells in temporal retina. In the present study we have attempted to falsify this hypothesis by investigating the effects of neonatal bilateral enucleation on the organization of callosal linkages in striate cortex of the rat. Once enucleated rats reached adulthood, we studied the mediolateral organization of callosal connections by placing small injections of different fluorescent tracers into different loci within medial and lateral striate cortex. The analysis of the distribution of retrogradely labeledcallosal cells indicated that connections from lateral portions of striate cortex were no longer organized in a reversed fashion, rather, they resembled the mirror image pattern normally found in the medial callosal region, i. e., injections at the 17/18a border produced labeled cells at the opposite 17/18a border, whereas injections into slightly more medial regions produced labeled cells in the opposite, mirror-symmetric location. In addition, we found that enucleation does not alter the organization of callosal linkages in medial portions of striate cortex. Thus, by showing that enucleation significantly changes the pattern of connections from lateral portions of striate cortex, the present study does not falsify, but rather strengthens the hypothesis that interhemispheric correlated activity driven from the temporal retinal crescent guides the normal development of reversed callosal linkages in lateral portions of rat striate cortex. Furthermore, the present study shows that, in the absence of the eyes, the pattern of callosal linkages in lateral portions of striate cortex resembles the mirror image pattern normally found only in medial striate cortex. © 1995 Wiley-Liss, Inc.     

Callosal connections of the posterior neocortex in normal-eyed, congenitally anophthalmic, and neonatally enucleated mice

Following multiple injections of horseradish peroxidase into the posterior neocortex of one hemisphere, we examined the distribution of retrogradely labeled cells and anterogradely labeled terminations in tangential and coronal sections through contralateral areas 17 and 18 in three groups of adult mice: normal-eyed (ZRDCT-n and C57Bl/6J strains), congenitally anophthalmic (ZRDCT-an strain), neonatally enucleated (ZRDCT-n strain). In agreement with previous studies, we observed that the pattern of callosal connections in areas 17 and 18 of normal-eyed mice contains the following features: (1) a dense band of callosal cells and terminations separating the interiors of areas 17 and 18, which have relatively few callosal connections, (2) a ring-like configuration anterolateral to area 17, (3) a region of dense labeling lateral to area 18, (4) a narrow band of labeling bridging the posterior portion of area 18, and (5) a region of labeling anteromedial to area 17. We find that all these features of the normal callosal pattern are recognizable in congenitally anophthalmic mice. Their presence in mice that never had eyes supports the hypothesis that central visual pathways can develop many aspects of their connectivity in the absence of input from the periphery. However, we also find that the details of certain features of the callosal pattern in congenitally eyeless mice often differ from those of the same features in normal-eyed mice, and that the between-animal variability in the appearance of these features is higher in eyeless mice. These latter findings indicate that the eyes are needed during normal development to fine-tune the pattern of callosal connections. Our results also reveal that the callosal pattern in neonatally enucleated mice does not differ significantly from that in congenitally anophthalmic mice, indicating that the period in which the eyes guide callosal development extends into postnatal life. While the present data do not delineate the time course of this period, the finding of similarly abnormal callosal patterns in congenitally anophthalmic and neonatally enucleated mice suggests that the eyes exert little if any influence prenatally. Finally, examination of coronal sections indicates that the laminar distribution of callosal connections develops normally in both groups of eyeless mice.     

Laminar distribution of the cells of origin of the descending tectofugal pathways in the pigeon (Columba livia)

Injections of trkiated proline and horseradish peroxidase were used to study cortical connections of striate cortex in owl monkeys. All cases resulted in label in area 18 (V-II) and the middle temporal visual area (MT). In most, but not all cases, label was also present in the dorsomedial visual area (DM). The connections between striate cortex and each of these three visual areas were retinotopic and reciprocal. Projections to V-II, MT, and DM were concentrated in layers IV and III, while pyramidal cells in layers III, V, and VI of these areas projected back to striate cortex.     

Patterns of connections from the striate cortex to cortical visual areas in superior temporal sulcus of macaque and middle temporal gyrus of owl monkey

Double label-emulsion autoradiography has been used in rhesus monkeys to study cortico-cortical connections, arising from two separate sites of the striate cortex, and going in the same hemisphere into the cortex in the medial part of the posterior (or ventral) bank of the superior temporal sulcus (area STS). The projections from a single cortical locus are segregated into several patches of labeled terminals in layers III-IV, which form bands in consecutive sections. Fields of distribution of these bands from two sites of area 17 may completely overlap, when they arise from neighboring regions, or may partially overlap, when they arise from more separated regions. The latter show a retinotopical order in these connections and, consequently, of area STS itself: central visual field is lateral and ventral in STS to peripheral visual field, and upper visual field is ventral and medial to lower visual field. Connections from area 17 to area MT in owl monkey were analyzed in a comparative study. Results show that these connections are also highly segregated into bands, and that those arising in separate, but neighboring regions of 17, may greatly overlap in MT. The retinotopy of these connections was not studied, however. The similar divergent distribution of area 17 connections to STS and MT, and the similarity of the topography of STS connections and of the known retinotopy of area MT, are considered as elements of homologies between these areas. Functional implications of these divergent patterns of connections to these motion-sensitive areas (Zeki, '74), are considered.     

Optically imaged maps of orientation preference in primary visual cortex of cats and ferrets

Feature maps in the cerebral cortex constitute orderly representations of response features created within the cortex; an example is the mapping of orientation-selective neurons in visual cortex. We have compared the properties of orientation maps in area 17 of cats and ferrets, obtained by optical imaging of intrinsic signals. Orientation maps in both species contain a quasi-periodic distribution of iso-orientation domains that are organized into a lattice of pinwheels. However, the spatial density of orientation domains and of pinwheels in ferret area 17 is nearly twice that in cat area 17. The ferret map also contains more discontinuities, or fractures, where orientation changes abruptly. The size of orientation domains scales with interdomain spacing, so that the ratio of the two is approximately the same in both species. Consistent with this finding, the orientation tuning width of individual pixels is similar in the two. The magnitude of orientation preference, however, is much lower in ferret compared to cat. The greater incidence of fractures in ferret appears to be due to proportionately greater overlap between domains of different orientations, particularly along fracture lines that link pinwheel centers. We hypothesize that a key determinant of orientation maps, the relationship between orientation domain size and spacing, expresses an anatomical link between sizes of thalamocortical arbors and horizontal intracortical connections in area 17. J. Comp. Neurol. 387:358–370, 1997. © 1997 Wiley-Liss, Inc.     

Intrinsic laminar lattice connections in primate visual cortex

Intracortical injections of horseradish peroxidase (HRP) reveal a system of periodically organized intrinsic connections in primate striate cortex. In layers 2 and 3 these connections form a reticular or latticelike pattern, extending for about 1.5-2.0 mm around an injection. This connectional lattice is composed of HRP-labeled walls (350-450 microns apart Saimiri and about 500-600 microns in macaque) surrounding unlabeled central lacunae. Within the lattice walls there are regularly arranged punctate loci of particularly dense HRP label, appearing as isolated patches as the lattice wall labeling thins further from the injection site. A periodic organization has also been demonstrated for the intrinsic connections in layer 4B, which are apparently in register with the supragranular periodicities, although separated from these by a thin unlabeled region. The 4B lattice is particularly prominent in squirrel monkey, extending for 2-3 mm from an injection. In both layers, these intrinsic connections are demonstrated by orthogradely and retrogradely transported HRP and seem to reflect a system of neurons with long horizontal axon collaterals, presumably with arborizations at regularly spaced intervals. The intrinsic connectional lattice in layers 2 and 3 resembles the repetitive array of cytochrome oxidase activity in these layers; but despite similarities of dimension and pattern, the two systems do not appear identical. In primate, as previously described in tree shrews (Rockland et al., '82), the HRP-labeled anatomical connections resemble the pattern of 2-deoxyglucose accumulation resulting from stimulation with oriented lines, although the functional importance of these connections remains obscure.     

Corticocortical projections to area 1 in squirrel monkeys (Saimiri sciureus)

Cortical area 1 is a non‐primary somatosensory area in the primate anterior parietal cortex that is critical to tactile discrimination. The corticocortical projections to area 1 in squirrel monkeys were determined by placing multiple injections of anatomical tracers into separate body part representations defined by multiunit microelectrode mapping in area 1. The pattern of labeled cells in the cortex indicated that area 1 has strong intrinsic connections within each body part representation and has inputs from somatotopically matched regions of areas 3b, 3a, 2 and 5. Somatosensory areas in the lateral sulcus, including the second somatosensory area (S2), the parietal ventral area (PV), and the presumptive parietal rostral (PR) and ventral somatosensory (VS) areas, also project to area 1. Topographically organized projections to area 1 also came from the primary motor cortex (M1), the dorsal and ventral premotor areas (PMd and PMv), and the supplementary motor area (SMA). Labeled cells were also found in cingulate motor and sensory areas on the medial wall of the hemisphere. Previous studies revealed a similar pattern of projections to area 1 in Old World macaque monkeys, suggesting a pattern of cortical inputs to area 1 that is common across anthropoid primates.     

Model-based analysis of excitatory lateral connections in the visual cortex

Excitatory lateral connections within the primary visual cortex are thought to link neurons with similar receptive field properties. Here we studied whether this rule can predict the distribution of excitatory connections in relation to cortical location and orientation preference in the cat visual cortex. To this end, we obtained orientation maps of areas 17 or 18 using optical imaging and injected anatomical tracers into these regions. The distribution of labeled axonal boutons originating from large populations of excitatory neurons was then analyzed and compared with that of individual pyramidal or spiny stellate cells. We demonstrate that the connection patterns of populations of nearby neurons can be reasonably predicted by Gaussian and von Mises distributions as a function of cortical location and orientation, respectively. The connections were best described by superposition of two components: a spatially extended, orientation-specific and a local, orientation-invariant component. We then fitted the same model to the connections of single cells. The composite pattern of nine excitatory neurons (obtained from seven different animals) was consistent with the assumptions of the model. However, model fits to single cell axonal connections were often poorer and their estimated spatial and orientation tuning functions were highly variable. We conclude that the intrinsic excitatory network is biased to similar cortical locations and orientations but it is composed of neurons showing significant deviations from the population connectivity rule.