Interconnections of the auditory cortical fields of the cat with the cingulate and parahippocampal cortices

Summary The interconnections of the auditory cortex with the parahippocampal and cingulate cortices were studied in the cat. Injections of the anterograde and retrograde tracer WGA-HRP were performed, in different cats (n = 9), in electrophysiologically identified auditory cortical fields. Injections in the posterior zone of the auditory cortex (PAF or at the PAF/AI border) labeled neurons and axonal terminal fields in the cingulate gyrus, mainly in the ventral bank of the splenial sulcus (a region that can be considered as an extension of the cytoarchitectonic area Cg), and posteriorly in the retrosplenial area. Labeling was also present in area 35 of the perirhinal cortex, but it was sparser than in the cingulate gyrus. Following WGA-HRP injection in All, no labeling was found in the cingulate gyrus, but a few neurons and terminals were labeled in area 35. In contrast, no or very sparse labeling was observed in the cingulate and perirhinal cortices after WGA-HRP injections in the anterior zone of the auditory cortex (AI or AAF). A WGA-HRP injection in the cingulate gyrus labeled neurons in the posterior zone of the auditory cortex, between the posterior ectosylvian and the posterior suprasylvian sulci, but none was found more anteriorly in regions corresponding to AI, AAF and AII. The present data indicate the existence of preferential interconnections between the posterior auditory cortex and the limbic system (cingulate and parahippocampal cortices). This specialization of posterior auditory cortical areas can be related to previous observations indicating that the anterior and posterior regions of the auditory cortex differ from each other by their response properties to sounds and their pattern of connectivity with the auditory thalamus and the claustrum.Abbreviations AAF anterior auditory cortical field - aes anterior ectosylvian sulcus - AI primary auditory cortical field - AII secondary auditory cortical field - ALLS anterior-lateral lateral suprasylvian visual area - BF best frequency - C cerebral cortex - CC corpus callosum - CIN cingulate cortex - CL claustrum - DLS dorsal lateral suprasylvian visual area - DP dorsoposterior auditory area - E entorhinal cortex - IC inferior colliculus - LGN lateral geniculate nucleus - LV pars lateralis of the ventral division of the MGB - LVe lateral ventricule - MGB medial geniculate body - OT optic tract - OV pars ovoidea of the ventral division of the MGB - PAF posterior auditory cortical field - pes posterior ectosylvian sulcus - PLLS posterior-lateral lateral suprasylvian visual area - PS posterior suprasylvian visual area - PU putamen - RE reticular complex of thalamus - rs rhinal sulcus - SC superior colliculus - SS suprasylvian sulcus - T temporal auditory cortical field - TMB tetramethylbenzidine - VBX ventrobasal complex of thalamus, external nucleus - VL pars ventrolateralis of the ventral division of the MGB - VLS ventrolateral suprasylvian visual area - VPAF ventroposterior auditory cortical field - WGA-HRP wheat germ agglutinin labeled with horseradish peroxidase - wm white matter     

Connections of the anterior ectosylvian visual area (AEV)

Summary We have previously described a visual area situated in the cortex surrounding the deep infolding of the anterior ectosylvian sulcus of the cat (Mucke et al. 1982). Using orthograde and retrograde transport methods we now report anatomical evidence that this anterior ectosylvian visual area (AEV) is connected with a substantial number of both cortical and subcortical regions. The connections between AEV and other cortical areas are reciprocal and, at least in part, topographically organized: the rostral AEV is connected with the bottom region of the presylvian sulcus, the lower bank of the cruciate sulcus, the rostral part of the ventral bank of the splenial sulcus, the rostral portion of the lateral suprasylvian visual area (LS) and the lateral bank of the posterior rhinal sulcus; the caudal AEV is connected with the bottom region of the presylvian sulcus, the caudal part of LS, the ventral part of area 20 and the lateral bank of the posterior rhinal sulcus. Subcortically, AEV has reciprocal connections with the ventral medial thalamic nucleus (VM), with the medial part of the lateralis posterior nucleus (LPm), as well as with the lateralis medialis-suprageniculate nuclear (LM-Sg) complex. These connections are also topographically organized with more rostral parts of AEV being related to more ventral portions of the LPm and LM-Sg complex. AEV also projects to the caudate nucleus, the putamen, the lateral amygdaloid nucleus, the superior colliculus, and the pontine nuclei. It is concluded that AEV is a visual association area which functionally relates the visual with both the motor and the limbic system and that it might play a role in the animal's orienting and alerting behavior.Abbreviations Ac aqueductus cerebri - AEs anterior ectosylvian sulcus - ALLS anterolateral lateral suprasylvian area - AMLS anteromedial lateral suprasylvian area - ASs anterior suprasylvian sulcus - Cd caudate nucleus - CL central lateral nucleus - Cl claustrum - Cos coronal sulcus - Crs cruciate sulcus - DLS dorsal lateral suprasylvian area - GI stratum griseum intermediale - GP stratum griseum profundum - IC inferior colliculus - LAm lateral amygdaloid nucleus - LGNd dorsal nucleus of lateral geniculate body - LGNv ventral nucleus of lateral geniculate body - Llc nucleus lateralis intermedius, pars caudalis - LM nucleus lateralis medialis - LPl nucleus lateralis posterior, pars lateralis - LPm nucleus lateralis posterior, pars medialis - Ls lateral sulcus - MD nucleus mediodorsalis - MG medial geniculate body - MSs middle suprasylvian sulcus - Ndl nucleus dorsolateralis pontis - Nl nucleus lateralis pontis - Np nucleus peduncularis pontis - Npm nucleus paramedianus pontis - Nrt nucleus reticularis tegmenti pontis - Nv nucleus ventralis pontis - Ped cerebral peduncle - PEs posterior ectosylvian sulcus - Pg periaqueductal gray - PLLS posterolateral lateral suprasylvian area - PMLS posteromedial lateral suprasylvian area - PSs presylvian sulcus - Pul pulvinar - Put putamen - R red nucleus - Sg suprageniculate nucleus - SN substantia nigra - Sps splenial sulcus - Syls sylvian sulcus - T trapezoid body - VA ventral anterior nucleus - VL ventral lateral nucleus - VLS ventral lateral suprasylvian area - VM ventral medial nucleus - VPL ventral posterolateral nucleus - VPM ventral posteromedial nucleus Sponsored by Max-Planck-Society during part of the studySponsored by Thyssen FoundationSponsored by Alexander von Humboldt-Foundation     

The association connexions of the suprasylvian fringe (SF) and other areas of the cat auditory cortex

The association connexions of the peri-auditory (SF, Ea and INS) and auditory (AI, AII and Ep) areas of the cat cortex were studied in silver impregnated material of 32 experiments with cortical lesions. The cortex of the lateral bank of the rostral part of the middle suprasylvian sulcus (SF) sends many fibres to AI and to the insular cortex (INS), and has scanty projections upon AII and Ep. In addition, it sends fibres to the visual area 17 as well as to the ventral bank of the medial part of the cruciate sulcus. It receives fibres from the three auditory areas AI, AII and Ep, as well as from Ea and INS. The dorsal part of the anterior ectosylvian gyrus (Ea) projects upon SF, AI, and AII. Ea sends few fibres to Ep, and receives relatively dense projections from AI and AII. The anterior sylvian gyrus (INS) projects heavily upon AII as well as upon the superficial part of SF. It sends a few fibres also to Ep. INS receives heavy projections from AII and relatively lighter connections from SF, AI and Ep. The three auditory areas AI, AII and Ep are strongly mutually interconnected. AI and Ep have scanty projections upon the visual area 19, and AI also to the lateral suprasylvian visual area, as well as upon the ventral bank of the medial cruciate sulcus. Correlations of the association connexions with the functions of each area are discussed.     

Physiologic and anatomic investigation of a visual cortical area situated in the ventral bank of the anterior ectosylvian sulcus of the cat

Summary In this paper a cortical area is described that covers approximately the posterior two-thirds of the ventral bank of the anterior ectosylvian sulcus of the cat and is called anterior ectosylvian visual area (AEV).In cats anesthetized with a combination of N₂O and barbiturate we explored this area by recording extracellularly the responses of AEV neurons to visual and electric stimulation as well as by injecting HRP into physiologically verified points. AEV neurons were found to be highly sensitive to small light stimuli moving rapidly in a particular direction through their large receptive fields. The properties of 74 neurons were quantitatively analyzed. Increasing the length of the stimulus within the receptive field to more than 2 deg strongly inhibited the responses, whereas increasing the speed of the stimulus movement up to 72–120 deg/s enhanced the neuronal responsiveness. Although the majority of neurons responded to a wide range of possible directions, one clearly preferred direction could usually be found for each neuron. There was a predominance of preferred directions toward the contralateral hemifield. Anatomic and electrophysiologic connectivity studies showed that AEV receives its main afferent inputs from the lateral suprasylvian visual area (LS) and from the tecto-recipient zone of the nucleus lateralis posterior (LP)-pulvinar complex.Although these studies suggested some topographical organization within the projection from LS to AEV, the large receptive fields in AEV, the great majority of which included the central area, did not reveal a clear retinotopic order. It is concluded that AEV is a specific visual area and that functionally the extrageniculate inputs predominate.Abbreviations AEs anterior ectosylvian sulcus - ALLS anterolateral lateral suprasylvian area - AMLS anteromedial lateral suprasylvian area; - Cl Claustrum - DLS dorsal lateral suprasylvian area - LGNd dorsal nucleus of lateral geniculate body - LGNv ventral nucleus of lateral geniculate body - LM nucleus lateralis medialis - LP1a nucleus lateralis posterior, pars lateralis - LPm nucleus lateralis posterior, pars medialis - Ls lateral sulcus - MGmc magnocellular division of medial geniculate body - MGpc parvocellular division of medial geniculate body - MSs middle suprasylvian sulcus - NP nucleus posterior of Rioch - PLLS postero-lateral lateral suprasylvian area - PLs posterolateral sulcus - PMLS posteromedial lateral suprasylvian area - Pu putamen - Pul pulvinar - Sg suprageniculate nucleus - VLS ventral lateral suprasylvian area Sponsored by Max-Planck-Gesellschaft and IBRODept. of Anatomy, School of Medicine, Iwate Medical University, Morioka 020, JapanSponsored by Alexander von Humboldt-FoundationDept. of Physiology, University Medical School, Szeged, Hungary     

Cortical control of sound localization in the cat: unilateral cooling deactivation of 19 cerebral areas

We examined the ability of mature cats to accurately orient to, and approach, an acoustic stimulus during unilateral reversible cooling deactivation of primary auditory cortex (AI) or 1 of 18 other cerebral loci. After attending to a central visual stimulus, the cats learned to orient to a 100-ms broad-band, white-noise stimulus emitted from a central speaker or 1 of 12 peripheral sites (at 15 degrees intervals) positioned along the horizontal plane. Twenty-eight cats had two to six cryoloops implanted over multiple cerebral loci. Within auditory cortex, unilateral deactivation of AI, the posterior auditory field (PAF) or the anterior ectosylvian sulcus (AES) resulted in orienting deficits throughout the contralateral field. However, unilateral deactivation of the anterior auditory field, the second auditory cortex, or the ventroposterior auditory field resulted in no deficits on the orienting task. In multisensory cortex, unilateral deactivation of neither ventral or dorsal posterior ectosylvian cortices nor anterior or posterior area 7 resulted in any deficits. No deficits were identified during unilateral cooling of the five visual regions flanking auditory or multisensory cortices: posterior or anterior ii suprasylvian sulcus, posterior suprasylvian sulcus or dorsal or ventral posterior suprasylvian gyrus. In motor cortex, we identified contralateral orienting deficits during unilateral cooling of lateral area 5 (5L) or medial area 6 (6m) but not medial area 5 or lateral area 6. In a control visual-orienting task, areas 5L and 6m also yielded deficits to visual stimuli presented in the contralateral field. Thus the sound-localization deficits identified during unilateral deactivation of area 5L or 6m were not unimodal and are most likely the result of motor rather than perceptual impairments. Overall, three regions in auditory cortex (AI, PAF, AES) are critical for accurate sound localization as assessed by orienting.     

Auditory projections to extrastriate visual cortex: connectional basis for multisensory processing in ‘unimodal’ visual neurons

Neurophysiological studies have recently documented multisensory properties in ‘unimodal’ visual neurons of the cat posterolateral lateral suprasylvian (PLLS) cortex, a retinotopically organized area involved in visual motion processing. In this extrastriate visual area, a region has been identified where both visual and auditory stimuli were independently effective in activating neurons (bimodal zone), as well as a second region where visually-evoked activity was significantly facilitated by concurrent auditory stimulation but was unaffected by auditory stimulation alone (subthreshold multisensory region). Given their different distributions, the possible corticocortical connectivity underlying these distinct forms of crossmodal convergence was examined using biotinylated dextran amine (BDA) tracer methods in 21 adult cats. The auditory cortical areas examined included the anterior auditory field (AAF), primary auditory cortex (AI), dorsal zone (DZ), secondary auditory cortex (AII), field of the rostral suprasylvian sulcus (FRS), field anterior ectosylvian sulcus (FAES) and the posterior auditory field (PAF). Of these regions, the DZ, AI, AII, and FAES were found to project to the both the bimodal zone and the subthreshold region of the PLLS. This convergence of crossmodal inputs to the PLLS suggests not only that complex auditory information has access to this region but also that these connections provide the substrate for the different forms (bimodal versus subthreshold) of multisensory processing which may facilitate its functional role in visual motion processing.     

Sensory modality distribution in the anterior ectosylvian cortex (AEC) of cats

Modality specificity of neuronal responses to visual, somesthetic and auditory stimuli was investigated in the anterior ectosylvian cortex (AEC) of cats, using single-unit recording techniques. Seven classes of neurons were found, and according to their responsiveness to sensory stimuli regrouped into three categories: unimodal, bimodal and trimodal. Unimodal cells that responded to only one of the three stimulus modalities formed 59% of the units; 30.2% were bimodal, in that they showed a clear increase of neuronal discharges to two of the three stimulus types; 10.8% were defined as trimodal because they responded to all three stimulus modalities. Although the different categories of cells were intermingled within the AEC, indicating a certain degree of overlap between sensory modalities, some clustering of cell types was nonetheless evident. Thus, the somatosensory responsive cells were mainly located in the anterior two-thirds of the dorsal bank of the anterior ectosylvian sulcus. Visually responsive cells were concentrated on the ventral bank of the sulcus, whereas neurons with an auditory response occupied the banks and fundus of the posterior three-quarters of the sulcus. The histological distribution and physiological properties of AEC neurons suggest that this cortical region is a higher-order associative area whose function may be to integrate information from different sensory modalities.     

Thalamic projections to the posterior sylvian and posterior ectosylvian gyri of the sheep brain, revealed with the retrograde transport of horseradish peroxidase

Summary The auditory area of the sheep cerebral cortex was studied on the basis of its afferents from the medial geniculate nucleus, traced with the horseradish peroxidase retrograde transport method. The results show that the medial geniculate nucleus projects only to the anterior parts of the posterior ectosylvian gyrus and the posterior sylvian gyrus. A small area of the posterior ectosylvian gyrus receives afferents exclusively from the ventral part of the medial geniculate nucleus, while the anterior part of the posterior sylvian gyrus receives also afferents from the posterior nucleus of the thalamus and the pulvinar. In addition, it was found that the medial part of the medial geniculate nucleus projects in a sparse way to the auditory cortex. The middle part of the posterior ectosylvian gyrus receives afferents from the posterior nucleus of the thalamus, the suprageniculate nucleus and the pulvinar, while the posterior part of the posterior ectosylvian gyrus together with the posteriormost part of the posterior sylvian gyrus receive afferents from the pulvinar. Finally, the area located between the anterior and the posteriormost part of the posterior sylvian gyrus receives afferents from both the posterior nucleus of the thalamus and the pulvinar.Abbreviations Ad nucleus anterior dorsalis - Am nucleus anterior medialis - Av nucleus anterior ventralis - BCI nucleus of the brachium colliculi inferioris - bci brachium colliculi inferioris - Cg substantia grisea centralis - ci capsula interna - Cm nucleus centralis medialis - EC sulcus ectomarginalis - EN sulcus entomarginalis - Ep epiphysis - ES sulcus ectosylvius - fd columna fornicis descendens - FS fissura sylvia - Hl nucleus habenularis lateralis - Hm nucleus habenularis medialis - Iv nucleus interventralis - Ld nucleus lateralis dorsalis - LGN nucleus geniculatus lateralis - LGNd nucleus geniculatus lateralis, pars dorsalis - LGNv nucleus geniculatus lateralis, pars ventralis - lme lamina medullaris thalami externa - Lp nucleus lateralis posterior - Lt nucleus lateralis thalami - MA sulcus marginalis - Md nucleus medialis dorsalis - MGN nucleus geniculatus medialis - MGNd nucleus geniculatus medialis, pars dorsalis - MGNm nucleus geniculatus medialis, pars magnocellularis - MGNv nucleus geniculatus medialis, pars ventralis - MIN medial interlaminar nucleus - mt fasciculus mamillothalamicus - ml lemniscus medialis - Mv nucleus medialis ventralis - ot tractus opticus - p putamen - pc pedunculus cerebri - Pl nucleus paralemniscalis - Po nucleus posterior - Pp nucleus paraventricularis posterior - Pta nucleus praetectalis anterior - Ptp nucleus praetectalis posterior - Pul pulvinar - R nucleus ruber - rf fasciculus retroflexus - Rh nucleus rhomboidalis - RH sulcus rhinalis lateralis - Rt nucleus reticularis thalami - Sg nucleus suprageniculatus - SN substantia nigra - SP sulcus cinguli - SS sulcus suprasylvius - Sth nucleus subthalamicus - Va nucleus ventralis anterior - Vl ventrolateral nuclear complex - Vll pars lateralis of the ventrolateral nuclear complex - Vm nucleus ventralis medialis - Vp nucleus ventralis posterior - Vpl nucleus ventralis posterior, pars lateralis - Vpm nucleus ventralis posterior, pars medialis - W Wernicke's field     

Quantitative analyses of principal and secondary compound parieto-occipital feedback pathways in cat

The purpose of our study was to quantify the magnitude of principal and secondary pathways emanating from the middle suprasylvian (MS) region of visuoparietal cortex and terminating in area 18 of primary visual cortex. These pathways transmit feedback signals from visuoparietal cortex to primary visual cortex. (1) WGA-HRP was injected into area 18 to identify inputs from visual structures. In terms of numbers of neurons, feedback projections to area 18 from MS sulcal cortex (areas PMLS, AMLS and PLLS) comprise 26% of inputs from all visual structures. Of these neurons, between 21% and 34.9% are located in upper layers 2–4 and the dominant numbers are located in deep layers 5 and 6. Areas 17 (11.8%) and 19 (11.2%) provide more modest cortical inputs, and another eight areas provide a combined total of 4.3% of inputs. The sum of neurons in all subcompartments of the lateral geniculate nucleus (LGN) accounts for another 34.8% of the input to area 18, whereas inputs from the lateral division of the lateral-posterior nucleus (LPl) account for the final 11.9%. (2) Injection of tritiated-(³H)-amino acids into MS sulcal cortex revealed substantial direct projections from MS cortex that terminated in all layers of area 18, but with a markedly lower density in layer 4. Projections from MS cortex to both areas 17 and 19 are of similar density and characteristics, whereas those to other cortical targets have very low densities. Quantification also revealed minor-to-modest axon projections to all components of LGN and a massive projection throughout the LP-Pul complex. (3) Superposition of the labeled terminal and cell fields identified secondary, compound feedback pathways from MS cortex to area 18. The largest secondary pathway is massive and it includes the LPl nucleus. Much more modest secondary pathways include areas 17 and 19, and LGN. The relative magnitudes of the secondary pathways suggest that the one through LPl exerts a major influence on area 18, whereas the others exert more modest or minor influences. MS cortex in the contralateral hemisphere also innervates area 18 directly. These data are important for interpreting the impact of deactivating feedback projections from visuoparietal cortex on occipital cortex.Abbreviations A layer A of LGN - A1 layer A1 of LGN - ALLS anterolateral visual area of the lateral suprasylvian sulcus (Palmer et al. 1978) - AMLS anteromedial visual area of the lateral suprasylvian sulcus (Palmer et al. 1978) - Aud auditory cortex of the middle ectosylvian gyrus - CC corpus callosum - Cg cingulate gyrus - Cm magnocellular layers of LGN - Cp parvocellular layers of LGN - LGN dorsal lateral geniculate nucleus - LP lateral posterior nucleus - LPl lateral division of the lateral posterior nucleus - LPm medial division of the lateral posterior nucleus (Graybiel and Berson 1980, Berson and Graybiel 1978; Raczkowski and Rosenquist 1983) - MIN medial interlaminar nucleus subdivision of LGN - MS cortex bounding the middle suprasylvian sulcus (areas AMLS, ALLS, PMLS, and PLLS) - OR optic radiation - PE posterior ectosylvian visual cortex - PLLS posterolateral visual area of the lateral suprasylvian sulcus (Palmer et al. 1978) - PMLS posteromedial visual area of the lateral suprasylvian sulcus (Palmer et al. 1978) - Pul pulvinar nucleus - SVA splenial visual area - V1 primary visual cortex - V2 secondary visual cortex - V3 third visual area - V5/MT fifth visual area/middle temporal area - WGA-HRP wheat germ agglutinin conjugated to horseradish peroxidase - Wing wing of LGN - 7 area 7 - 17 area 17 - 18 area 18 - 19 area 19     

Topographical and laminar distribution of cortical input to the monkey entorhinal cortex

Hippocampal formation plays a prominent role in episodic memory formation and consolidation. It is likely that episodic memory representations are constructed from cortical information that is mostly funnelled through the entorhinal cortex to the hippocampus. The entorhinal cortex returns processed information to the neocortex. Retrograde tracing studies have shown that neocortical afferents to the entorhinal cortex originate almost exclusively in polymodal association cortical areas. However, the use of retrograde studies does not address the question of the laminar and topographical distribution of cortical projections within the entorhinal cortex. We examined material from 60 Macaca fascicularis monkeys in which cortical deposits of either (3)H-amino acids or biotinylated dextran-amine as anterograde tracers were made into different cortical areas (the frontal, cingulate, temporal and parietal cortices). The various cortical inputs to the entorhinal cortex present a heterogeneous topographical distribution. Some projections terminate throughout the entorhinal cortex (afferents from medial area 13 and posterior parahippocampal cortex), while others have more limited termination, with emphasis either rostrally (lateral orbitofrontal cortex, agranular insular cortex, anterior cingulate cortex, perirhinal cortex, unimodal visual association cortex), intermediate (upper bank of the superior temporal sulcus, unimodal auditory association cortex) or caudally (parietal and retrosplenial cortices). Many of these inputs overlap, particularly within the rostrolateral portion of the entorhinal cortex. Some projections were directed mainly to superficial layers (I-III) while others were heavier to deep layers (V-VI) although areas of dense projections typically spanned all layers. A primary report will provide a detailed analysis of the regional and laminar organization of these projections. Here we provide a general overview of these projections in relation to the known neuroanatomy of the entorhinal cortex.     

Afferent connections to the pontine nuclei from the cortex of the anterior ectosylvian sulcus in the cat

Following injections of horseradish peroxidase conjugated with wheatgerm agglutinin (HRP-WGA) in different sectors of the cortex of the anterior ectosylvian sulcus (SEsA), anterograde labeling was observed in the pontine nuclei (PN) of the cat. Labeled fibers were identified in a wide area which covers the entire rostro-caudal extent of the PN. The various sectors of the SEsA, which differ in their associative, cortico-cortical connections with the somatosensory, auditory and visual cortices, also were shown to differ in their projection patterns to the PN. These corticopontine projections of the SEsA were compared to those from the modality specific areas.     

Corticotectal and corticothalamic efferent projections of SIV somatosensory cortex in cat

Substantial corticotectal (and corticothalamic) projections from the cortex of the anterior ectosylvian sulcus (AES) were demonstrated in the cat using the axonal transport methods of autoradiography and horseradish peroxidase. The corticotectal projection arises nearly exclusively from medium-large pyramidal cells in lamina V. One of the densest projecting areas of the AES is the rostral aspect of its superior bank, where a fourth somatotopic representation (SIV) has recently been demonstrated. It terminates in the intermediate and deep laminae of the superior colliculus, where somatic cells are located. The pathway is bilateral but much heavier ipsilaterally than contralaterally. In contrast to the substantial corticotectal projection from SIV and adjacent tissue, there was no unequivocal evidence for a corticotectal projection from traditional somatosensory cortex SI-SIII. This finding, that somatosensory projections to the cat superior colliculus arise from an area outside of SI-SIII, was unexpected on the basis of what is known about visual corticotectal projections. However, it is consistent with the patterns of other cortical projections that terminate in the intermediate and deep laminae of this structure and with the absence of demonstrable corticotectal influences from SI to SIII in this animal. These data are in contrast to demonstrations by other investigators that there is a corticotectal projection from SI cortex in rodents. Apparently there is a fundamental species difference in the organization of descending somatosensory pathways. A corticothalamic projection of the AES was also observed. This descending projection appeared to form a shell of labeled cells and fibers around the ventrobasal complex, but unequivocal terminal labeling within the ventrobasal complex could not be demonstrated. Dense terminal labeling was apparent in the posterior group of thalamic nuclei (PO) where thalamocortical afferents to the AES originate.     

Organization of a fourth somatosensory area of cortex in cat

The organization of sensory representations in the cortex of the anterior ectosylvian sulcus (AES) of the cat was investigated using single-unit recording techniques. Somatic, auditory, and visual cells were found in the AES but were partially segregated. Somatic cells were concentrated in the rostral two-thirds of the sulcus, auditory cells were found in the caudal third, and visual cells were distributed along the fundus. A distinct, heretofore unknown, somatotopic representation of the body surface was observed in the AES and was designated SIV. The representation of the body in SIV extends along a rostrocaudal axis and the entire somatotopic map is inverted, with the head rostral and the hindquarters caudal. The representation of the paws extends over the lip of the sulcus to abut the paw representations in SII, and the SIV-SII boundary is marked by a reversal in the sequence of receptive fields along the AEG-AES. The SIV representation (SII) on the crown of the anterior ectosylvian gyrus (AEG). The somatotopic map in SII was found to extend further lateral on the AEG than shown by some investigations and it contains a double representation of the limbs: a large representation with the limbs having the opposite orientation to and abutting the SIV map and a smaller representation located more medial on the AEG and extending into the suprasylvian sulcus. The presence of this double representation may help to explain previous discrepancies regarding the overall orientation of the body in SII.     

Somatosensory and multisensory properties of the medial bank of the ferret rostral suprasylvian sulcus

In ferret cortex, the rostral portion of the suprasylvian sulcus separates primary somatosensory cortex (SI) from the anterior auditory fields. The boundary of the SI extends to this sulcus, but the adjoining medial sulcal bank has been described as “unresponsive.” Given its location between the representations of two different sensory modalities, it seems possible that the medial bank of the rostral suprasylvian sulcus (MRSS) might be multisensory in nature and contains neurons responsive to stimuli not examined by previous studies. The aim of this investigation was to determine if the MRSS contained tactile, auditory and/or multisensory neurons and to evaluate if its anatomical connections were consistent with these properties. The MRSS was found to be primarily responsive to low-threshold cutaneous stimulation, with regions of the head, neck and upper trunk represented somatotopically that were primarily connected with the SI face representation. Unlike the adjoining SI, the MRSS exhibited a different cytoarchitecture, its cutaneous representation was largely bilateral, and it contained a mixture of somatosensory, auditory and multisensory neurons. Despite the presence of multisensory neurons, however, auditory inputs exerted only modest effects on tactile processing in MRSS neurons and showed no influence on the averaged population response. These results identify the MRSS as a distinct, higher order somatosensory region as well as demonstrate that an area containing multisensory neurons may not necessarily exhibit activity indicative of multisensory processing at the population level.     

Deoxyglucose uptake in the ferret auditory cortex

Histological methods and 2-deoxyglucose autoradiography were used in an attempt at finding distinguishing characteristics that would permit the clear definition of different auditory areas on the ectosylvian gyrus. This region was studied in both coronal and flattened tangential sections. In tangential sections a crescent-shaped region of high deoxyglucose uptake was identified. The centre of this crescent was in the position of the primary auditory area on the middle ectosylvian gyrus. The ventro-anterior arm of the crescent was on the surface of the anterior ectosylvian gyrus and the ventro-posterior arm on the posterior ectosylvian gyrus. All three parts of the crescent appear to have an auditory function, because ablating the inferior colliculus or inserting a contralateral earplug reduced their deoxyglucose uptake. This was shown by using two separately distinguishable forms of 2-deoxyglucose, incorporating the ¹⁸F and ¹⁴C isotopes. In addition, another area of high deoxyglucose activity was identified in the ventral wall of the suprasylvian sulcus, which seems to correspond to the anterior auditory field. These four areas with high deoxyglucose uptake also have high levels of succinate dehydrogenase activity and moderately high densities of myelinated fibres. Succinate dehydrogenase histochemistry provides a simple method for identifying auditory cortical areas and should be of use in future physiological studies. These results provide evidence that the ferret has four separate auditory areas with relatively high metabolic and functional activity. Received: 8 August 1996 / Accepted: 15 May 1997     

Corticocortical projections to the prefrontal cortex in the rhesus monkey investigated with horseradish peroxidase techniques

The corticocortical afferents innervating the prefrontal cortex in the monkey were studied by means of the retrograde axonal transport of horseradish peroxidase. After injection of small amounts (0.3-0.5 microliter) of this enzyme into various parts of the prefrontal cortex, many labeled neurons (mostly pyramids of 15-25 microns in diameter) were found in various cortical regions of the ipsilateral hemisphere. A small part of the prefrontal cortex received fibers from other parts of the same cortex. For example, area 8 receives many fibers from both the rostral part of area 9 and a small area adjacent to the inferior branch of the arcuate sulcus. On the other hand, area 9 in the inferior prefrontal convexity receives fibers from localized parts of areas 8 and 9 in the dorsolateral convexity as well as from area 6. It is also apparent that association connections from the dorsolateral to the inferior convexity are stronger than those going in the opposite direction. The prefrontal afferents from other cortical regions include many fibers originating from the posterior association cortex as well as some fibers arising in the cingulate and orbital gyri. The prefrontal cortex does not receive direct corticocortical fibers from the motor and "primary" sensory cortices. There is a topographic pattern in the prefrontal projections from the cortical walls (STs area) surrounding the superior temporal sulcus. Thus, the caudal half of the STs area projects to area 8 and a small adjacent part of area 9. The dorsal wall of the rostral half of the STs area projects to areas 9-12, the fundus to the inferior convexity, and the ventral wall only to the caudal part of the convexity. Projections from the circumjacent association cortex of the STs area to the prefrontal cortex as well as to the STs area are likewise found to be topographically organized. This suggests that certain parts of the posterior association cortex projecting to particular areas of the prefrontal cortex, also send fibers to those parts of the STs area which project to the same prefrontal areas.     

Cortical projections of the anterior thalamic nuclei in the cat

Summary Unilateral stereotaxic lesions were made in the anterior thalamic nuclei of the cat, and the ensuing terminal degeneration traced to the medial cortex by the methods of Nauta-Gygax and Fink-Heimer. The anterodorsal nucleus projects to the retrosplenial, postsubicular and presubicular areas. These projections appear to be organized in the dorsoventral direction. The posterior portion of the retrosplenial area receives no fibers from the anterodorsal nucleus. Fibers from this nucleus are distributed largely in layer I and in layer III and the deep portion of layer II of the posterior limbic cortex. The anteroventral nucleus sends fibers to the cingular area and parts of the retrosplenial, postsubicular and presubicular areas. These projections appear to be organized in a topical manner mediolaterally. When the lesion involves the parvocellular part of the nucleus, degeneration spreads to the lower lip, bank and fundus of the splenial sulcus. There appears to be an anteroposterior organization in the cortical projections of the anteroventral nucleus. Fibers from the anteroventral nucleus are distributed most profusely in layers IV and III and in the superficial portion of layer I of the posterior limbic cortex. The anteromedial nucleus sends fine fibers to the anterior limbic region and to the cingular, retrosplenial, postsubicular and presubicular areas. The cortical projections of the anteromedial nucleus appear to be topographically organized in the dorsoventral direction. Fibers from the anteromedial nucleus are distributed largely in cortical layers V and VI of the anterior and posterior limbic regions.Abbreviations used in Figures a anterior - AD anterodorsal nucleus - AM anteromedial nucleus - AMD dorsolateral part of anteromedial nucleus - AMV ventromedial part of anteromedial nucleus - AV anteroventral nucleus - AVM magnocellular part of anteroventral nucleus - AVP parvocellular part of anteroventral nucleus - CC corpus callosum - Cg cingular area - CM medial central nucleus - Il infralimbic area - LA anterior limbic region - LD dorsal lateral nucleus - MD dorsal medial nucleus - Of orbitofrontal region - p posterior - Pr presubicular area - Prag precentral agranular area - Ps postsubicular area - Pt paratenial nucleus - Pv anterior paraventricular nucleus - R reuniens nucleus - Rs retrosplenial area - Rt thalamic reticular nucleus - SC cruciate sulcus - SM stria medullaris - Sm submedial nucleus - SS splenial sulcus - VA ventral anterior nucleus - VL ventral lateral nucleus - VM ventral medial nucleus     

Somato-sensory paths to the second cortical projection area of the group I muscle afferents

1. Cats anaesthetized with chloralose and paralysed with Flaxedil were used. The projections of muscle, joint and skin afferents to the cortical fold hidden in the anterior suprasylvian sulcus were investigated with micro-electrode recording techniques.2. Electrical stimulation of Group I muscle afferents from the contralateral forelimb evoked a negative focal potential (latency 5 msec) in a locus of 1-2 mm diameter found in the lower bank of the fold. In one experiment a response to Group I muscle afferents from the contralateral hind limb was observed. The Group I potentials disappeared after sectioning of the dorsal columns at C3.3. Groups II and III muscle afferents, low threshold skin afferents and joint afferents also evoked potentials in the Group I locus. It was concluded that the joint afferents originated mainly in the Ruffini endings of the joint capsule.4. Groups II and III muscle afferents, low threshold skin and low threshold joint afferents projected to the upper bank of the suprasylvian fold. A certain somatotopic arrangement was observed.5. The possibility of connexions between the cortex of the anterior suprasylvian fold and the primary somato-sensory projection areas was discussed, as well as the organization of the loci in the fold in terms of cell colonies with different properties.     

Cortical afferents and efferents of monkey postarcuate area: an anatomical and electrophysiological study

Summary A study has been made of the corticocortical efferent and afferent connections of the posterior bank of the arcuate sulcus in the macaque monkey. The distribution of efferent projections to the primary motor cortex (MI) was studied by injecting three different fluorescent retrograde tracers into separate regions of MI. The resultant labeling showed a discrete and topographically organized projection: neurons lying below the inferior limb of the arcuate sulcus project into the MI face area, while neurons located in the posterior bank of the inferior limb of the arcuate sulcus and in the arcuate spur region project into the MI hand area. These findings were confirmed electrophysiologically by demonstrating that postarcuate neurons could only be activated antidromically by stimulation within restricted regions of MI. HRP injections within postarcuate cortex indicated that afferents to this region arise from a number of cortical areas. However, the largest numbers of labeled neurons were found in the posterior parietal cortex (area 7b; PF) and in the secondary somatosensory region (SII). Neurons in both 7b (PF) and SII could be antidromically activated by postarcuate stimulation. It was further shown that stimulation of area 7b (PF) gives rise to short-latency synaptic responses in postarcuate neurons, including some neurons with identified projections to MI. The results are discussed in relation to the possible function of the postarcuate region of the premotor cortex in the sensory guidance of movement.     

Prefrontal cortex of the cat: Paucity of afferent projections from the parietal cortex

Summary Twenty-one cat brains with cortical injections of horseradish peroxidase resulting in labelled cells in the thalamic mediodorsal nucleus (MD) were screened for afferent projections from the parietal cortex. Contrary to expectation, nearly the whole prefrontal cortex (PFC) situated around the frontal pole was free of parietal afferents, while a small area in the anterior sylvian gyrus (orbito-insular subregion of PFC) consistently received afferents from the parietal cortex. The few afferents projecting to the cortex around the frontal pole originated exclusively from the convexity of the suprasylvian gyrus, while the great majority of the parietal neurons projecting to the anterior sylvian gyrus was situated within the fundus of the suprasylvian sulcus. While the main regions of the prefrontal cortex of the rhesus monkey receive a substantial projection from the parietal lobe, whereas the main regions of the cat's prefrontal cortex are free of afferents from the parietal cortex, possible differences in the parieto-prefrontal organization of both species are discussed. Furthermore, differences between the orbito-insular subregion and the rest of the PFC are emphasized.This study was carried out mainly at the University of Konstanz.Dr. B. Petrovi-Mini was a visiting scientist at the University of Konstanz. Research was supported in part by grant Ma 795 from the Deutsche Forschungsgemeinschaft (DFG)