Post-rolandic cortical projections of the superior temporal sulcus in the rhesus monkey.

The efferent connections of different cytoarchitectonic areas of the superior temporal sulcus (STS) in the rhesus monkey with parieto-temporo-occipital cortex were investigated using autoradiographic methods. Four rostral-to-caudal subdivisions of cortex (area TPO) in the upper bank of the STS have distinct projection patterns. Rostral sectors (areas TPO-1 and -2) project to the rostral superior temporal gyrus (areas Ts1, Ts2, and Ts3), insula of the Sylvian fissure, and parahippocampal gyrus (perirhinal and prorhinal cortexes, areas TF, TH, and TL); caudal sectors (TPO-3 and -4) project to the caudal superior temporal gyrus (areas paAlt and Tpt), supratemporal plane (area paAc), circular sulcus of the Sylvian fissure (area reIt), as well as medial paralimbic (areas 23, 24, and retrosplenial cortex) and extrastriate (areas 18 and 19) cortexes. Area TPO-1 does not project to the parietal lobe; area TPO-2 projects to the inferior parietal lobule; area TPO-3 to the lower bank of the intraparietal sulcus (IPS) (area POa); and area TPO-4 to medial parietal cortex (area PGm). Vision-related cortex (area TEa) in the rostral lower bank of the STS sends fibers to the rostral inferotemporal region (areas TE1, -2, and -3) and parahippocampal gyrus (perirhinal cortex, areas TF and TL). Visual zones in the caudal lower bank and depth of the sulcus (area OAa, or MT and FST) project to the caudal inferotemporal region (areas TE3 and TEO), lateral preoccipital region (area V4), and lower bank of the IPS (area POa). A zone in the rostral depth of the STS (area IPa) projects to the rostral inferotemporal region, parahippocampal gyrus, insula of the Sylvian fissure, parietal operculum, and lower rim of the IPS (area PG). STS projections to parieto-temporo-occipital cortex have "feedforward," "feedbackward," and "side-to-side" laminar patterns of termination similar to those of other cortical sensory systems. The differential connectivity supports the cytoarchitectonic parcellation of the STS and suggests functional heterogeneity.     

Striatal connections of the parietal association cortices in rhesus monkeys

The corticostriatal connections of the parietal association cortices were examined by the autoradiographic technique in rhesus monkeys. The results show that the rostral portion of the superior parietal lobule projects predominantly to the dorsal portion of the putamen, whereas the caudal portion of the superior parietal lobule and the cortex of the upper bank of the intraparietal sulcus have connections with the caudate nucleus as well as with the dorsal portion of the putamen. The medial parietal convexity cortex projects strongly to the caudate nucleus, and has less extensive projections to the putamen. In contrast, the medial parietal cortex within the caudal portion of the cingulate sulcus projects predominantly to the dorsal portion of the putamen, and has only minor connections with the caudate nucleus. The rostral portion of the inferior parietal lobule projects mainly to the ventral sector of the putamen, and has only minor connections with the caudate nucleus. The middle portion of the inferior parietal lobule has sizable projections to both the putamen and the caudate nucleus. The caudal portion of the inferior parietal lobule as well as the lower bank of the intraparietal sulcus project predominantly to the caudate nucleus, and have relatively minor connections with the putamen. The cortex of the parietal opercular region also shows a specific pattern of corticostriatal projections. Whereas the rostral portion projects exclusively to the ventral sector of the putamen, the caudal portion has connections to the caudate nucleus as well. Thus, it seems that parietostriatal projections show a differential topographic distribution; within both the superior and the inferior parietal region, as one progresses from rostral to caudal, there is a corresponding shift in the predominance of projections from the putamen to the caudate nucleus. In addition, with regard to the projections to the putamen, the superior parietal lobule is related mainly to the dorsal portion, and the inferior parietal lobule to the ventral portion. The striatal projections of the cortex of the caudal portion of the cingulate gyrus (corresponding in part to the supplementary sensory area) and of the rostral parietal opercular region (corresponding in part to the second somatosensory area) are directed almost exclusively to the dorsal and ventral sectors of the putamen, respectively. This pattern resembles that of the primary somatosensory cortex. The results are discussed with regard to the overall architectonic organization of the posterior parietal region. Possible functional aspects of parietostriatal connectivity are considered in the light of physiological and behavioral studies. © 1993 Wiley-Liss, Inc.     

Efferent association pathways originating in the caudal prefrontal cortex in the macaque monkey

The efferent association fibers from the caudal part of the prefrontal cortex to posterior cortical areas course via several pathways: the three components of the superior longitudinal fasciculus (SLF I, SLF II, and SLF III), the arcuate fasciculus (AF), the fronto-occipital fasciculus (FOF), the cingulate fasciculus (CING F), and the extreme capsule (Extm C). Fibers from area 8Av course via FOF and SLF II, merging in the white matter of the inferior parietal lobule (IPL) and terminating in the caudal intraparietal sulcus (IPS). A group of these fibers turns ventrally to terminate in the caudal superior temporal sulcus (STS). Fibers from the rostral part of area 8Ad course via FOF and SLF II to the IPS and IPL and via the AF to the caudal superior temporal gyrus and STS. Some fibers from the rostral part of area 8Ad are conveyed to the medial parieto-occipital region via FOF, to the STS via Extm C, and to the caudal cingulate gyrus via CING F. Fibers from area 8B travel via SLF I to the supplementary motor area and area 31 in the caudal dorsal cingulate region and via the CING F to cingulate areas 24 and 23 and the cingulate motor areas. Fibers from area 9/46d course via SLF I to the superior parietal lobule and medial parieto-occipital region, via SLF II to the IPL. Fibers from area 9/46v travel via SLF III to the rostral IPL and the frontoparietal opercular region and via the CING F to the cingulate gyrus.     

Parietal,temporal, and occipita projections to cortex of the superior temporal sulcus in the rhesus monkey: A retrograde tracer study

The afferent cortical connections of individual cytoarchitectonic areas within the superior temporal sucus (STS) of the rhesus monkey were studied by retrograde tracer techniques, including double tracer experiments. Rostral superior temporal polysensory (STP) cortex (area TPO-1) receives input from the rostral superior temporal gyrus (STG), cortex of the circular sulcus, and parahippocampal gyrus (PHG) (areas 35, TF, and TL). Mid-STP cortex (areas TPO-2 and -3) has input from the mid-STG, cortex of the mid-circular sulcus, caudal inferior parietal lodule (IPL), cingulate gyrus (areas, 23, 24, retrosplenial cortex), and mid-PHG (areas 28, TF, TH, and TL). Caudal STP cortex (area TPO-4) has afferent connections with the caudal STG, cortex of the cauda insula and caudal circular sulcus, caudal IPL, lower bank of the intraparietal sulcus (IPS), medial parietal lobe, cingulate gyrus, and mid- and caudal PHG (areas TF, TH, TL; prostriate area). The most rostral cortex of the lower bank of the STS (areasTEa and TEm), a presumed visual association area, receives input from the rostal inferotemporal (IT) region; more cauda portions of areas TEa and TEm have afferent connections with the caudal IT region, PHG, preoccipital gyrus, and cortex of the lower bank of the IPS. © 1994 Wiley-Liss, Inc.

1 This article is a US Government work and, as such, is in the public domain in the United States of America.

     

Cortical connections of the inferior arcuate sulcus cortex in the macaque brain

Injections of the retrograde/anterograde tracers Wheat Germ Agglutinin-Horseradish peroxidase (WGA-HRP) into the cortex along the banks of the inferior limb of the arcuate sulcus in the cortex of 4 macaque monkeys (Macaca fascicularis) were used to investigate its cortico-cortical connections. All injections produced transported label within the sulcus principalis, the ventral lateral prefrontal cortex, the anterior cingulate sulcus and the dorsal insular cortex. The distribution of label within each of these areas differed slightly depending on the injection site. Injections along the caudal bank of the inferior arcuate sulcus label premotor, supplementary motor, and precentral motor areas but produce relatively sparse prefrontal labeling. Posteriorly label is transported to the inferior parietal cortex and the dorsal opercular bank of the Sylvian fissure. Injections along the rostral bank of the sulcus do not label motor areas but produce labeling in dorsal, lateral and orbital prefrontal areas, and in cortex along the ventral bank of the superior branch of the arcuate sulcus. Posteriorly label is transported to cortical areas in the superior temporal gyrus including the dorsal bank of the superior temporal sulcus. The more dorsal rostral bank injection produced both superior temporal and some sparse inferior parietal labeling and the more ventral rostral bank injection produced extensive superior temporal labeling but no parietal labeling. No labeling was ever seen in cortex ventral to the fundus of the superior temporal sulcus. Although other auditory recipient prefrontal areas have been reported, this is the first demonstration of a region chiefly devoted to auditory connections within the ventral frontal cortex. Its adjacency to areas associated with vocal muscle movement, and its connections to midline cortical areas associated with vocal functions in both primates and humans may provide important clues to the organization of Broca's language area.     

Inferior parietal lobule. Divergent architectonic asymmetries in the human brain

Architectonic parcellation of the parietal lobes of eight human brains, with special attention paid to the inferior parietal lobule, resulted in a map that bore a close relationship to previous maps and took into consideration modern data on physiology and connections. Two general parietal zones were distinguished, one above and the other below the intraparietal sulcus, similar to the dorsal-ventral distinction suggested for the frontal lobe. Five areas were recognized in the inferior parietal lobule, of which areas parietal areas EG (PEG), G (PG), and occipitoparietal G (OPG) were in the angular gyrus. A lateralization toward the right was found for area PEG, an area structurally similar to the visually related cortices of the posterior superior parietal region. A lateralization toward the left was found for area PG, but only in brains with a larger left planum temporale. The asymmetry in area PG seemed to be linked to other asymmetries present in language areas, whereas the right-sided area PEG preponderance showed no relation to the language asymmetries.     

Fiber pathways and cortical connections of preoccipital areas in rhesus monkeys

An understanding of visual function at the cerebral cortical level requires detailed knowledge of anatomical connectivity. Cortical association pathways and terminations of preoccipital visual areas were investigated in rhesus monkeys by using the autoradiographic tracing technique. Medial and adjacent dorsomedial preoccipital regions project via the occipitofrontal fascicle to the frontal lobe (dorsal area 6, and areas 8Ad, 8B, and 46); via the dorsal portion of the superior longitudinal fascicle (SLF) to dorsal area 6, area 9, and the supplementary motor area; and via the cingulate fascicle to area 24. In addition, medial and dorsomedial preoccipital areas send projections to parietal (areas PGm, PEa, PG‐Opt, and POa) and superior temporal (areas MST and MT) regions. In contrast, connections from the dorsolateral, annectant, and ventral preoccipital regions are conveyed via the inferior longitudinal fascicle (ILF) to the parietal lobe (areas POa and IPd), superior temporal sulcus (areas MT, MST, FST, V4t, and IPa), inferotemporal region (areas TEO and TE1–TE3), and parahippocampal gyrus (areas TF, TH, and TL). The central‐lateral preoccipital region projects via an ILF‐SLF pathway to frontal area 8Av. The preoccipital areas also have caudal connections to occipital areas V1, V2, and V3. Finally, preoccipital regions are interconnected via different intrinsic pathways. These findings provide further insight into the nature of preoccipital fiber pathways and the connectional organization of the visual system. J. Comp. Neurol. 518:3725–3751, 2010. © 2010 Wiley‐Liss, Inc.     

Cortical connections of the visuomotor parietooccipital area V6Ad of the macaque monkey

Area V6A, a functionally defined region in the anterior bank of the parietooccipital sulcus, has been subdivided into dorsal and ventral cytoarchitectonic fields (V6Ad and V6Av). The aim of this study was to define the cortical connections of the cytoarchitectonic field V6Ad. Retrograde and bidirectional neuronal tracers were injected into the dorsal part of the anterior bank of parietooccipital sulcus of seven macaque monkeys (Macaca fascicularis). The limits of injection sites were compared with those of cytoarchitectonic fields. The major connections of V6Ad were with areas of the superior parietal lobule. The densest labeling was observed in the medial intraparietal area (MIP). Areas PEc, PGm, and V6Av were also strongly connected. Labeled cells were found in medial parietal area 31; in cingulate area 23; in the anterior (AIP), ventral (VIP), and lateral (LIP) intraparietal areas; in the inferior parietal lobule (fields Opt and PG); and in the medial superior temporal area (MST). In the frontal lobe, the main projection originated from F2, although labeled cells were also found in F7 and area 46. Preliminary results obtained from injections in nearby areas PEc and V6Av revealed connections different from those of V6Ad. In agreement with functional data, the strong connections with areas where arm‐reaching activity is represented suggest that V6Ad is part of a parietofrontal circuit involved in the control of prehension, and connections with AIP specifically support an involvement in the control of grasping. Connections with areas LIP and Opt are likely related to the oculomotor activities observed in V6Ad. J. Comp. Neurol. 513:622–642, 2009. © 2009 Wiley‐Liss, Inc.     

Chemoarchitectonics and corticocortical terminations within the superior temporal sulcus of the rhesus monkey: Evidence for subdivisions of superior temporal polysensory cortex

Cortex of the upper bank of the superior temporal sulcus (STS) in macaque monkeys, termed the superior temporal polysensory (STP) region, corresponds largely to architectonic area TPO and is connectionally distinct from adjacent visual areas. To investigate whether or not the STP region contains separate subdivisions, immunostaining for parvalbumin and neurofilament protein (using the SMI-32 antibody) was compared with patterns of corticocortical terminations in the STS. Chemoarchitectonic results provided evidence for three caudal-to rostral subdivisions: TPOc, TPOi, and TPOr. Area TPOc was characterized by patchy staining for parvalbumin and SMI-32 in cortical layers IV/III and III, respectively. Area TPOi had more uniform chemoarchitectonic staining, whereas area TPOr had a thicker layer IV than TPOi. The connectional results showed prefrontal cortex in the location of the frontal eye fields (area8) and dorsal area 46 projected in a columnar pattern to all cortical layers of area TPOc, to layer IV of TPOi, and in a columanr fashion, with a moderate increase in density in layer IV, to TPOr. In TPOc, columns of frontal connections showed a peridicity similar to that of the SMI-32 staining. The caudal inferior parietal lobule (area 7a) and superior temporal gyrus projected to each subdivision of area TPO, displaying either panlaminar or fourth-layer terminations. In addition to STP cortex, parvalbumin and SMI-32 immunostaining allowed identification of caudal visual areas of the STS, including MT, MST, FST, and V4t. These areas received first and sixth-layer projections from prefrontal cortex and area 7a. © 1995 Wiley-Liss, Inc.     

The cortico-cortical connections within the parieto-temporal lobe of area PG,7a, in the monkey

After injections of HRP into area PG(7a) labelled cells have been found in architectonic areas OA, PE, the cingulate and retrosplenial areas medial to area PG; posteriorly areas MST, OA (V4), V2, V3 and the cortex in the walls and floor of the superior temporal sulcus have also been labelled. Small injections placed in PG have resulted in different parts of these areas being labelled, suggesting that these cortico-cortical connections are well organized and raising the possibility of an ordered representation of the visual field in PG. It is suggested that the vertical meridian is around the boundary and the horizontal meridian passes antero-posteriorly across about the middle of its medio-lateral extent; the central part of the visual field is in the depths of the intraparietal sulcus, and the periphery is on the surface of the inferior parietal lobule and in the anterior wall of the upper part of the superior temporal sulcus. The lower visual field is medial and the upper field is lateral.     

Corticothalamic connections of the posterior parietal cortex in the rhesus monkey

Corticothalamic connections of posterior parietal regions were studied in the rhesus monkey by using the autoradiographic technique. Our observations indicate that the rostral superior parietal lobule (SPL) is connected with the ventroposterolateral (VPL) thalamic nucleus. In addition, whereas the rostral SPL is connected with the ventrolateral (VL) and lateral posterior (LP) thalamic nuclei, the rostral IPL has connections with the ventroposteroinferior (VPI), ventroposteromedial parvicellular (VPMpc), and suprageniculate (SG) nuclei as well as the VL nucleus. The caudal SPL and the midportion of IPL show projections mainly to the lateral posterior (LP) and oral pulvinar (PO) nuclei, respectively. These areas also have minor projections to the medial pulvinar (PM) nucleus. Finally, the medial SPL and the caudal IPL project heavily to the PM nucleus, dorsally and ventrally, respectively. In addition, the medial SPL has some connections with the LP nucleus, whereas the caudal IPL has projections to the lateral dorsal (LD) nucleus. Furthermore, the caudal and medial SPL and the caudal IPL regions have additional projections to the reticular and intralaminar nuclei-the caudal SPL predominantly to the reticular, and the caudal IPL mainly to the intralaminar nuclei. These results indicate that the rostral-to-caudal flow of cortical connectivity within the superior and inferior parietal lobules is paralleled by a rostral-to-caudal progression of thalamic connectivity. That is, rostral parietal association cortices project primarily to modality-specific thalamic nuclei, whereas more caudal regions project most strongly to associative thalamic nuclei.     

Converging visual and somatic sensory cortical input to the intraparietal sulcus of the rhesus monkey

A cyto- and myeloarchitectonic study reveals the presence of a distinct cortical zone ("area POa") in the lower bank of the intraparietal sulcus of the rhesus monkey. Using both autoradiographic and silver impregnation techniques, an analysis of cortical connections shows two overlapping projections to this sulcal zone. These come from (1) the middle portion of the preoccipital gyrus (area OA) and (2) the rostral inferior parietal lobule (area PF).     

Connections of the parietal lobe

The efferent connections of the posterior parietal cortex were studied in rhesus monkeys subjected to selective lesions of the superior and inferior parietal lobules, which correspond approximately to Brodmann's areas 5 and 7, respectively.

Following ablations of either the superior or inferior parietal lobule, axon degeneration, stained with the Nauta and Fink-Heimer methods, was traced into the extreme, external, and internal capsules, and into the cerebral peduncle. This degeneration extended into the ipsilateral insular cortex, cingulate gyrus, prefrontal and premotor cortices, and the precentral and postcentral gyri. In addition to these connections, the superior lobule sends fibers to the ipsilateral inferior parietal lobule and superior temporal gyrus, and via the corpus callosum to the contralateral superior and inferior parietal lobules, whereas the inferior parietal lobule sends fibers to the ipsilateral superior parietal lobule and to the contralateral superior and inferior parietal lobules. A prominent fiber system to the ipsilateral temporal lobe degenerates following lesions in the inferior parietal lobule (area 7); in such cases fiber degeneration appears in the superior, middle and inferior temporal convolutions, and in the fusiform and parahippocampal gyri.

Both lobules evidently project to the claustrum and body of the caudate nucleus. Both, moreover, have massive efferent connections with the dorsal two-thirds of the putamen. By contrast, no evidence of projections from the parietal cortex to the globus pallidus was found in any of the cases studied.

A further subcortical projection from the posterior parietal cortex involves the nucleus reticularis thalami and the nucleus lateralis posterior thalami. The inferior lobule projects directly to the nucleus lateralis dorsalis and to the mediodorsal region of the nucleus lateralis posterior that closely adjoins two thalamic cell groups: the n. lateralis dorsalis and the intralaminar nucleus centralis lateralis. The superior parietal lobule, by contrast, projects massively to a ventrolateral district of the nucleus lateralis posterior.

Parietosubthalamic connections could be traced from areas 5 and 7 to the zona incerta and fields H₂ and H of Forel, but evidence for terminal connections with the n. subthalamicus (Luys) could not be foud.

Both areas 5 and 7 project massively to the pretectal area and the deeper layers of the superior colliculus. This parieto-mesencephalic connection is amplified by a fiber connection from the inferior parietal lobule (area 7) to the lateral, densocellular region of the circumaqueductal gray matter. No evidence of parietal corticonigral fibers connections was found. Finally, both parietal lobules were found to project to the pontine nuclei.

Speculations regarding the associative functions of the parietal lobules at the cortical and subcortical levels are presented, with particular emphasis upon the possible significance of the projections from the inferior parietal lobule to insular, cingulate and temporal regions of the cortex.     

Afferent and efferent projections of the inferior area 6 in the macaque monkey

The rostral part of the agranular frontal cortex (area 6) can be subdivided on the basis of its cytoarchitecture, enzymatic properties, and connections into two large sectors: a superior region, lying medial to the spur of the arcuate sulcus, and an inferior region, lying lateral to it. In this study we traced the afferent and efferent connections of the inferior region of area 6 by injecting small amounts of wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP) and fluorescent tracers (fast blue and diamidino yellow) into restricted parts of inferior area 6 and in physiologically determined fields of area 4. There is an ordered topographic pattern of connections between inferior area 6 and area 4. The region near the spur of the arcuate sulcus (hand field) projects to the area 4 hand field while the lateral part of inferior area 6 (mouth field) is connected with the corresponding field in area 4. The organization of the connections between the two fields is, however, different. The hand fields in area 6 and 4 have direct reciprocal projections, whereas the mouth field in the postarcuate cortex relays information to area 4 via a zone intermediate between the arcuate and the central sulcus. This zone corresponds to the cytochrome oxidase area F4 (Matelli, Luppino, and Rizzolatti: Behav. Brain Res. 18: 125-137, '85). The inferior area 6 also has topographically organized connections with the supplementary motor area. The inferior area 6 receives and sends fibers to a series of discrete cortical areas located in the lower cortical moiety (Sanides: The Structure and Function of the Nervous Tissue, Vol. 5. New York: Academic Press, pp 329-453, '72). These areas that form a broad ring around the central sulcus are the ventral bank of the principal sulcus and the adjacent area 46, the precentral operculum (PrOC), area SII (Jones and Burton: J. Comp. Neurol. 168:197-248, '76), the parietal operculum, and the rostral part of the inferior parietal lobule including the lower bank of the intraparietal sulcus. Finally, the inferior area 6 has sparse but consistent connections with insular and cingulate cortices. The functional significance of this complex pattern of connections is discussed.     

The connections of area PG, 7a, with cortex in the parietal, occipital and temporal lobes of the monkey

The cortico-cortical connections of area PG, 7a, in the parietal, occipital and temporal lobes have been studied after injections of HRP in this area and in certain of the areas connected with it. After such injections in PG there are labelled cells in architectonic areas OA and PE (visual area PO), the cingulate and retrosplenial areas situated medial to PG; posteriorly labelled cells are present in OA, visual areas MST, MT, V2, V3, V4 and in the walls and floor of the lower part of the superior temporal sulcus. Injections in PE and V4 show that these connections are reciprocal. Small injections in PG result in cell labelling in different parts of the areas connected to PG, suggesting that the connections are well organized and that there may be an ordered representation of the visual field in PG. In the lower wall of the lower part of the superior temporal sulcus there is overlap of the two visual pathways in the cortex, that to the temporal lobe with that to the parietal lobe; and in a restricted part of this sulcus there is convergence and overlap of the sequences of cortico-cortical connections related to the visual, somatic and auditory sensory systems. There may be certain common principles in the sequences of cortical connections to the parietal and temporal lobes from the primary visual and somatic sensory areas; in both there are well organized hierarchical and parallel pathways, and both are related to the superior temporal sulcus and to the cingulate cortex.     

Cingulate cortex of the rhesus monkey: II. Cortical afferents

Cortical projections to subdivisions of the cingulate cortex in the rhesus monkey were analyzed with horseradish peroxidase and tritiated amino acid tracers. These projections were evaluated in terms of an expanded cytoarchitectural scheme in which areas 24 and 23 were divided into three ventrodorsal parts, i.e., areas 24a-c and 23a-c. Most cortical input to area 25 originated in the frontal lobe in lateral areas 46 and 9 and orbitofrontal areas 11 and 14. Area 25 also received afferents from cingulate areas 24b, 24c, and 23b, from rostral auditory association areas TS2 and TS3, from the subiculum and CA1 sector of the hippocampus, and from the lateral and accessory basal nuclei of the amygdala (LB and AB, respectively). Areas 24a and 24b received afferents from areas 25 and 23b of cingulate cortex, but most were from frontal and temporal cortices. These included the following areas: frontal areas 9, 11, 12, 13, and 46; temporal polar area TG as well as LB and AB; superior temporal sulcus area TPO; agranular insular cortex; posterior parahippocampal cortex including areas TF, TL, and TH and the subiculum. Autoradiographic cases indicated that area 24c received input from the insula, parietal areas PG and PGm, area TG of the temporal pole, and frontal areas 12 and 46. Additionally, caudal area 24 was the recipient of area PG input but not amygdalar afferents. It was also the primary site of areas TF, TL, and TH projections. The following projections were observed both to and within posterior cingulate cortex. Area 29a-c received inputs from area 46 of the frontal lobe and the subiculum and in turn it projected to area 30. Area 30 had afferents from the posterior parietal cortex (area Opt) and temporal area TF. Areas 23a and 23b received inputs mainly from frontal areas 46, 9, 11, and 14, parietal areas Opt and PGm, area TPO of superior temporal cortex, and areas TH, TL, and TF. Anterior cingulate areas 24a and 24b and posterior areas 29d and 30 projected to area 23. Finally, a rostromedial part of visual association area 19 also projected to area 23. The origin and termination of these connections were expressed in a number of different laminar patterns. Most corticocortical connections arose in layer III and to a lesser extent layer V, while others, e.g., those from the cortex of the superior temporal sulcus, had an equal density of cells in both layers III and V.(ABSTRACT TRUNCATED AT 400 WORDS)     

Anatomical investigation of projections from thalamus to posterior parietal cortex in the rhesus monkey: a WGA-HRP and fluorescent tracer study.

The parietothalamic projections have been shown to be heterogeneous and appear to be a reflection of the detailed architectonic parcellation of the parietal lobe. In the present study WGA-HRP injections were placed in the different subdivisions of the posterior parietal cortex of the rhesus monkey to determine whether a similarly complex pattern also exists in the thalamocortical pathway. Additionally, in an attempt to determine whether there is an intranuclear specificity of projections from individual thalamic nuclei to different subdivisions of the parietal lobe, multiple retrograde fluorescent tracers were injected into the rostral to caudal sectors of the parietal lobe of the same animal. Different subdivisions of the parietal lobe appear to receive different sets of thalamic input. Thus the superior parietal lobule (SPL) projections are derived from more lateral regions in the thalamus, arising predominantly from the lateral posterior (LP) and pulvinar oralis (PO) nuclei, with additional contributions from the pulvinar lateralis (PL) and pulvinar medialis (PM) nuclei. The inferior parietal lobule (IPL), by contrast, receives its projections from more medial thalamic regions, its main thalamic input originating from PM, and aided by LP, PL, and PO. Both the SPL and IPL also receive projections from the mediodorsal (MD), ventroposterior, ventrolateral, intralaminar, and limbic nuclei, albeit from different components within these nuclei. A topographical arrangement also exists in the thalamic projections to the rostral versus the caudal subdivisions of both the SPL and the IPL. Thus, in the SPL, the ventral posterolateral nucleus, pars oralis (VPLo), ventral lateral nucleus, pars oralis (VLo), and ventral lateral nucleus, pars medialis (VLm) project to rostral regions, whereas the PM and limbic nuclei, anteroventral (AV), and anteromedial (AM), project to area PGm on the medial convexity of the SPL. With respect to projections to the IPL, the ventral posteromedial (VPM) and PO nuclei project to rostral regions, whereas the limbic nuclei lateral dorsal (LD), AM and AV project only to the caudal most area, Opt. A rostrocaudal difference is reflected also within certain nuclei (LP, PO, and PM) that project to the SPL or IPL. Thus rostral parietal subdivisions receive projections from ventral regions within these thalamic nuclei, whereas caudal parietal afferents arise from the dorsal parts of these nuclei. Intervening cortical levels receive projections from intermediate positions within the nuclei. It therefore seems that the increasing architectonic and functional complexity as one moves from rostral to caudal in the SPL and IPL appear to be reflected in the thalamic afferents.(ABSTRACT TRUNCATED AT 400 WORDS)     

Afferent and efferent connections of the dorsolateral precentral gyrus (area 4, hand/arm region) in the macaque monkey, with comparisons to area 8

The afferent and efferent connections of the dorsolateral precentral gyrus, the primary motor cortex for control of the upper extremity, were studied by using the retrograde and anterograde capabilities of the horseradish peroxidase (HRP) technique in three adult macaque monkeys that had received HRP gel implants in this cortical region. Reciprocal corticocortical connections were observed primarily with the supplementary motor area (SMA) in medial premotor area 6 and dorsal bank of the cingulate sulcus, postarcuate area 6 cortex, dorsal cingulate cortex (area 24), superior parietal lobule (area 5, PE/PEa), and inferior parietal lobule (area 7b, PF/PFop, including the secondary somatosensory SII region). In these heavily labeled regions, the associational intrahemispheric afferents originated primarily from small and medium sized pyramidal cells in layer III, but also from layer V. The SMA projections were columnar in organization. Intrahemispheric afferents from contralateral homologous and nonhomologous frontal and cingulate cortices also originated predominantly from layer III, but the connections from contralateral area 4 were almost exclusively from layer III. The bilateral connections with premotor frontal area 6 and cingulate cortices were not observed with parietal regions; i.e., only ipsilateral intrahemispheric parietal corticocortical connections were observed. There were no significant connections with prearcuate area 8 or the granular frontal (prefrontal) cortex. Subcortical afferents originated primarily from the nucleus basalis of Meynert, dorsal claustrum, ventral lateral (VLo and VLc), area X, ventral posterolateral pars oralis (VPLo), central lateral and centromedian thalamic nuclei, lateral hypothalamus, pedunculopontine nucleus, locus ceruleus and subceruleus, and superior central and dorsal raphe nuclei. Lesser numbers of retrogradely labeled neurons were observed in the nucleus of the diagonal band, mediodorsal (MD), paracentral, and central superior lateral thalamic nuclei, nucleus limitans, nucleus annularis, and the mesencephalic and pontine reticular formation.(ABSTRACT TRUNCATED AT 400 WORDS)     

Further observations on corticofrontal connections in the rhesus monkey

The frontal lobe connections of the post-Rolandic sensory association areas are investigated. Our results indicate that the caudal portion of the superior temporal gyrus (area 22), the lateral peristriate belt (areas 18, 19), and the superior parietal lobule and the rostralmost portion of the inferior parietal lobule (areas 5 and 7), all project to periarcuate cortex, while the middle portion of area 22, caudal inferotemporal cortex (area 20), and the middle portion of the lower bank of the intraparietal sulcus, all have connections predominantly to prearcuate cortex. In contrast, rostral area 22 and the rostral inferotemporal cortex (area 21) project primarily to the orbital surface, and the middle portion of area 7 projects to the mid-principal sulcus. Those regions that projects to periacuate cortex are termed first association areas (AA1, VA1, SA1), those that project primarily to prearcuate cortex are designated second association areas (AA2, VA2, SA2), while those that project mainly to the orbital surface or the mid-principal sulcus are called third association areas (AA3, VA3, SA3). It was also found that the caudalmost portion of area 7 has a distinct projection pattern, connecting with the dorsal prearcuate cortex — areas 8B and 46. Additionally, it was observed that the connections from the association areas of different sensory modalities appear to overlap in specific areas of frontal cortex. Projections from the first association areas seem to converge in the periarcuate zone (bimodal overlap is noted between VA1 and AA1 in the arcuate concavity, and SA1 and VA1 dorsal to the arcuate sulcus), while those from the second association areas overlap in the ventral prearcuate cortex (area 46), where both bimodal and trimodal overlap is observed.