Auditory corticocortical interconnections in the cat: evidence for parallel and hierarchical arrangement of the auditory cortical areas

Summary The origin and laminar arrangement of the homolateral and callosal projections to the anterior (AAF), primary (AI), posterior (PAF) and secondary (AII) auditory cortical areas were studied in the cat by means of electrophysiological recording and WGA-HRP tracing techniques. The transcallosal projections to AAF, AI, PAF and AII were principally homotypic since the major source of input was their corresponding area in the contralateral cortex. Heterotypic transcallosal projections to AAF and AI were seen, originating from the contralateral AI and AAF, respectively. PAF received heterotypic commissural projections from the opposite ventroposterior auditory cortical field (VPAF). Heterotypic callosal inputs to AII were rare, originating from AAF and AI. The neurons of origin of the transcallosal connections were located mainly in layers II and III (70–92%), and less frequently in deep layers (V and VI, 8–30%). Single unit recordings provided evidence that both homotypic and heterotypic transcallosal projections connect corresponding frequency regions of the two hemispheres. The regional distribution of the anterogradely labeled terminals indicated that the homotypic and heterotypic auditory transcallosal projections are reciprocal. The present data suggest that the transcallosal auditory interconnections are segregated in 3 major parallel components (AAF-AI, PAF-VPAF and AII), maintaining a segregation between parallel functional channels already established for the thalamocortical auditory interconnections. For the intrahemispheric connections, the analysis of the retrograde tracing data revealed that AAF and AI receive projections from the homolateral cortical areas PAF, VPAF and AII, whose neurons of origin were located mainly in their deep (V and VI) cortical layers. The reciprocal interconnections between the homolateral AAF and AI did not show a preferential laminar arrangement since the neurons of origin were distributed almost evenly in both superficial (II and III) and deep (V and VI) cortical layers. On the contrary, PAF received inputs from the homolateral cortical fields AAF, AI, AII and VPAF, originating predominantly from their superficial (II and III) layers. The homolateral projections reaching AII originated mainly from the superficial layers of AAF and AI, but from the deep layers of VPAF and PAF. The laminar distribution of anterogradely labeled terminal fields, when they were dense enough for a confident identification, was systematically related to the laminar arrangement of neurons of origin of the reciprocal projection: a projection originating from deep layers was associated with a reciprocal projection terminating mainly in layer IV, whereas a projection originating from superficial layers was associated with a reciprocal projection terminating predominantly outside layer IV. This laminar distribution indicates that the homolateral auditory cortical interconnections have a feed-forward/feed-back organization, corresponding to a hierarchical arrangement of the auditory cortical areas, according to criteria previously established in the visual system of primates. The principal auditory cortical areas could be ranked into 4 distinct hierarchical levels. The tonotopically organized areas AAF and AI represent the lowest level. The second level corresponds to the non-tonotopically organized area AII. Higher, the tonotopically organized areas VPAF and PAF occupy the third and fourth hierarchical levels, respectively.Abbreviations AAF anterior auditory cortical area - AI primary auditory cortical area - AII secondary auditory cortical area - BF best frequency - C cerebral cortex - CA caudate nucleus - CL claustrum - D dorsal nucleus of the dorsal division of the MGB - ea anterior ectosylvian sulcus - ep posterior ectosylviansulcus - IC internal capsule - LGN lateral geniculate nucleus - LV pars lateralis of the ventral division of the MGB - LVe lateral ventricule - M pars magnocellularis of the medial division of the MGB - MGB medial geniculate body - MGBv ventral division of the MGB - OT optic tract - OV pars ovoidea of the ventral division of the MGB - PAF posterior auditory cortical area - PH parahippocampal cortex - PO lateral part of the posterior group of thalamic nuclei - PU putamen - RE reticular complex of thalamus - rs rhinal sulcus - SG suprageniculate nucleus of the dorsal division of the MGB - ss suprasylvian sulcus - TMB tetrametylbenzidine - VBX ventrobasal complex - VLa ventrolateral complex - VL ventro-lateral nucleus of the ventral division of the MGB - WGA-HRP wheat germ agglutinin conjugated to horse-radish peroxidase - WM white matter - VPAF ventro-posterior auditory cortical area     

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     

Overlapping projections to the amygdala and striatum from auditory processing areas of the thalamus and cortex.

The purpose of this study was to advance our understanding of the anatomical organization of sensory projections to the amygdala, and specifically to identify potential interactions within the amygdala between thalamic and cortical sensory projections of a single sensory modality. Thus, interconnections between the amygdala and acoustic processing areas of the thalamus and cortex were examined in the rat using WGA-HRP as an anterograde and a retrograde axonal tracer. Injections placed in medial aspects of the medial geniculate body (MGB) produced anterograde transport to the lateral nucleus of the amygdala and to adjacent areas of the striatum. Injections of primary auditory cortex (TE1) produced no transport to amygdala. In contrast, injections ventral to TE1 involving TE3 and perirhinal periallocortex (PRh) produced anterograde transport in the subcortical forebrain that was indistinguishable from that produced by the MGB injections. The TE3 and PRh injections also resulted in retrograde transport to primary auditory cortex and to MGB, thus confirming the involvement of these ventral cortical areas in auditory functions. Injections of the lateral nucleus of the amygdala resulted in retrograde transport back to the medial areas of MGB and to temporal cortical areas PRh, TE3, and the ventral most part of TE1. Thus, auditory processing regions of the thalamus and cortex give rise to overlapping (possibly convergent) projections to the lateral nucleus of the amygdala. These projections may allow diverse auditory signals to act on common ensembles of amygdaloid neurons and may therefore play a role in the integration of sensory messages leading to emotional reactions.     

Corticofugal modulation of the information processing in the auditory thalamus of the cat

Summary Single unit activity of 355 cells was recorded in the auditory thalamus of anesthetized cats before, during, and after the inactivation by cooling of the ipsilateral primary auditory cortex (AI). Most of the units (n = 288) showed similar functional characteristics of firing before and after the cryogenic blockade of AI. The spontaneous firing rate remained unchanged by cooling in 20% of the units and decreased in the majority of them (60%). In some regions, i.e. dorsal division of the medial geniculate body (MGB), lateral part of the posterior group of the thalamus, and auditory sector of the reticular nucleus of the thalamus, the maximum firing rate evoked by white noise bursts was generally affected by cooling in the same direction and to the same extent as the spontaneous activity. Units in the ventral division of MGB showed a characteristic increase of signal-to-noise ratio during cortical cooling. The corticofugal modulation led to the appearance or disappearance of the best frequency of tuning in 51 units and changed it by more than 0.5 octave in 34 units. The bandwidths of different response patterns to pure tones stimulation were used to define a set of functional properties. During cryogenic blockade of AI, two cortically modulated sub-populations of units were usually distinguished that exhibited changes for a given functional property. The complexity and diversity of the effects of cortical inactivation suggest that the corticothalamic projection may be the support for selective operations such as an adaptive filtering of the incoming acoustic signal at the thalamic level adjusted as a function of cortical activity.     

Neural correlates of gap detection in three auditory cortical fields in the Cat.

Neural correlates of gap detection in three auditory cortical fields in the cat. Mimimum detectable gaps in noise in humans are independent of the position of the gap, whereas in cat primary auditory cortex (AI) they are position dependent. The position dependence in other cortical areas is not known and may resolve this contrast. This study presents minimum detectable gap-in-noise values for which single-unit (SU), multiunit (MU) recordings and local field potentials (LFPs) show an onset response to the noise after the gap. The gap, which varied in duration between 5 and 70 ms, was preceded by a noise burst of either 5 ms (early gap) or 500 ms (late gap) duration. In 10 cats, simultaneous recordings were made with one electrode each in AI, anterior auditory field (AAF), and secondary auditory cortex (AII). In nine additional cats, two electrodes were inserted in AI and one in AAF. Minimum detectable gaps based on SU, MU, or LFP data in each cortical area were the same. In addition, very similar minimum early-gap values were found in all three areas (means, 36.1-41.7 ms). The minimum late-gap values were also similar in AI and AII (means, 11.1 and 11.7 ms), whereas AAF showed significantly larger minimum late-gap durations (mean 21.5 ms). For intensities >35 dB SPL, distributions of minimum early-gap durations in AAF and AII had modal values at approximately 45 ms. In AI, the distribution was more uniform. Distributions for minimum late-gap duration were skewed toward low values (mode at 5 ms), but high values (相似文献   

The auditory pathway in cat corpus callosum

The cortical auditory fields of the two hemispheres are interconnected via the corpus callosum. We have investigated the topographical arrangement of auditory callosal axons in the cat. Following circumscribed biocytin injections in the primary (AI), secondary (AII), anterior (AAF) and posterior (PAF) auditory fields, labelled axons have been found in the posterior two-thirds of the corpus callosum. Callosal axons labelled by small individual cortical injections did not form a tight bundle at the callosal midsagittal plane but spread over as much as one-third of the corpus callosum. Axons originating from different auditory fields were roughly topographically ordered, reflecting to some extent the rostro-caudal position of the field of origin. Axons from AAF crossed on average more rostrally than axons from AI; the latter crossed more rostrally than axons from PAF and AIL Callosal axons originating in a discrete part of the cortex travelled first in a relatively tight bundle to the telo-diencephalic junction and then dispersed progressively. In conclusion, the cat corpus callosum does not contain a sector reserved for auditory axons, nor a strictly topographically ordered auditory pathway. This observation is of relevance to neuropsychological and neuropathological observations in man.     

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.     

Sound localization during homotopic and heterotopic bilateral cooling deactivation of primary and nonprimary auditory cortical areas in the cat

Although the contributions of primary auditory cortex (AI) to sound localization have been extensively studied in a large number of mammals, little is known of the contributions of nonprimary auditory cortex to sound localization. Therefore the purpose of this study was to examine the contributions of both primary and all the recognized regions of acoustically responsive nonprimary auditory cortex to sound localization during both bilateral and unilateral reversible deactivation. The cats learned to make an orienting response (head movement and approach) to a 100-ms broad-band noise stimulus emitted from a central speaker or one of 12 peripheral sites (located in front of the animal, from left 90 degrees to right 90 degrees , at 15 degrees intervals) along the horizontal plane after attending to a central visual stimulus. Twenty-one cats had one or two bilateral pairs of cryoloops chronically implanted over one of ten regions of auditory cortex. We examined AI [which included the dorsal zone (DZ)], the three other tonotopic fields [anterior auditory field (AAF), posterior auditory field (PAF), ventral posterior auditory field (VPAF)], as well as six nontonotopic regions that included second auditory cortex (AII), the anterior ectosylvian sulcus (AES), the insular (IN) region, the temporal (T) region [which included the ventral auditory field (VAF)], the dorsal posterior ectosylvian (dPE) gyrus [which included the intermediate posterior ectosylvian (iPE) gyrus], and the ventral posterior ectosylvian (vPE) gyrus. In accord with earlier studies, unilateral deactivation of AI/DZ caused sound localization deficits in the contralateral field. Bilateral deactivation of AI/DZ resulted in bilateral sound localization deficits throughout the 180 degrees field examined. Of the three other tonotopically organized fields, only deactivation of PAF resulted in sound localization deficits. These deficits were virtually identical to the unilateral and bilateral deactivation results obtained during AI/DZ deactivation. Of the six nontonotopic regions examined, only deactivation of AES resulted in sound localization deficits in the contralateral hemifield during unilateral deactivation. Although bilateral deactivation of AI/DZ, PAF, or AES resulted in profound sound localization deficits throughout the entire field, the cats were generally able to orient toward the hemifield that contained the acoustic stimulus, but not accurately identify the location of the stimulus. Neither unilateral nor bilateral deactivation of areas AAF, VPAF, AII, IN, T, dPE, nor vPE had any effect on the sound localization task. Finally, bilateral heterotopic deactivations of AI/DZ, PAF, or AES yielded deficits that were as profound as bilateral homotopic cooling of any of these sites. The fact that deactivation of any one region (AI/DZ, PAF, or AES) was sufficient to produce a deficit indicated that normal function of all three regions was necessary for normal sound localization. Neither unilateral nor bilateral deactivation of AI/DZ, PAF, or AES affected the accurate localization of a visual target. The results suggest that hemispheric deactivations contribute independently to sound localization deficits.     

Contralateral connections of the dog's frontal association cortex

We studied the topography of contralateral connections of both prefrontal and premotor regions of the dog's frontal association cortex (FAC) by charting distributions of retrogradely labeled cells following unilateral HRP injections to various areas of this cortex. Generally, in the contralateral hemisphere the labeled cells were most numerous in the FAC areas localized homotopically to the injection sites, less numerous in FAC areas heterotopic to injections, and the least numerous in cortical areas situated outside the frontal lobe. The nonfrontal areas which project to the dorsal and ventral FAC differ from one another. Dorso-caudal parts of the cingular and insular areas, as well as the auditory, somatosensory and visual association cortices project to the dorsal FAC, while the ventro-rostral parts of the cingular and insular areas, together with the prepiriform and periamygdaloid areas of the olfactory cortex as well as the subcallosal area send their axons to the ventral FAC. Thus, the dorsal and ventral FAC areas are supplied by contralateral afferents originating from different cortical areas. Similar organization of ipsilateral FAC connections was described previously.     

Transcranial photo-inactivation of neural activities in the mouse auditory cortex

Flavoprotein fluorescence in the brain is intimately coupled with neuronal aerobic energy metabolism. If flavoproteins are photobleached, neural activities may be affected owing to dysfunction in aerobic energy metabolism in mitochondria. We tested this possibility in cortical slices from mice, and found that exposure to blue light (λ = 475 nm) derived from a 20 mW diode laser for 50 min suppresses trans-synaptic components of field potentials. This finding formed the basis of a transcranial photo-inactivation technique, that was used to investigate auditory signal transmission between the anterior auditory field (AAF) and the primary auditory cortex (AI) in anesthetized mice. Cortical responses in AAF and AI, elicited by 5 kHz tonal stimuli, were visualized using transcranial flavoprotein fluorescence imaging. After determining responsive areas in AAF and AI, the auditory cortex was exposed to the blue diode laser via the intact skull, while either AAF or AI was protected with a piece of carbon paper. Although the photo-inactivation of AI had no significant effect on the fluorescence responses in AAF, the photo-inactivation of AAF significantly reduced the fluorescence responses in AI, indicating the presence of auditory signal transmission from AAF to AI.     

Multiparametric auditory receptive field organization across five cortical fields in the albino rat

The auditory cortex of the rat is becoming an increasingly popular model system for studies of experience-dependent receptive field plasticity. However, the relative position of various fields within the auditory core and the receptive field organization within each field have yet to be fully described in the normative case. In this study, the macro- and micro-organizational features of the auditory cortex were studied in pentobarbital-anesthetized adult rats with a combination of physiological and anatomical methods. Dense microelectrode mapping procedures were used to identify the relative position of five tonotopically organized fields within the auditory core: primary auditory cortex (AI), the posterior auditory field (PAF), the anterior auditory field (AAF), the ventral auditory field (VAF), and the suprarhinal auditory field (SRAF). AI and AAF both featured short-latency, sharply tuned responses with predominantly monotonic intensity-response functions. SRAF and PAF were both characterized by longer-latency, broadly tuned responses. VAF directly abutted the ventral boundary of AI but was almost exclusively composed of low-threshold nonmonotonic intensity-tuned responses. Dual injection of retrograde tracers into AI and VAF was used to demonstrate that the sources of thalamic input from the medial geniculate body to each area were essentially nonoverlapping. An analysis of receptive field parameters beyond characteristic frequency revealed independent spatially ordered representations for features related to spectral tuning, intensity tuning, and onset response properties in AI, AAF, VAF, and SRAF. These data demonstrate that despite its greatly reduced physical scale, the rat auditory cortex features a surprising degree of organizational complexity and detail.     

Organization of corticopontocerebellar connections to the paramedian lobule in the cat

Summary To reveal the organization and relative magnitude of connections from various parts of the cerebral cortex to the cerebellar paramedian lobule via the pontine nuclei, horseradish peroxidase conjugated to wheat germ agglutinin was injected in the paramedian lobule in conjunction with injection of the same tracer in various parts of the cerebral cortex in 14 cats. Termination areas of cortical fibres (anterogradely labelled) and pontine neurons projecting to the paramedian lobule (retrogradely labelled) were carefully plotted in serial sections through the pons. On the average 89% of all labelled cells were found in the pontine nuclei contralateral to the cerebellar injection, 11% in the ipsilateral pontine nuclei. The highest degree of overlap between anterograde and retrograde labelling was found after injections in the posterior sigmoid gyrus (SmI), while less overlap was found after injections of the anterior sigmoid gyrus (MsI). Injections of the second somatosensory area (SmII) and the parietal association cortex (areas 5 and 7) gave moderate degrees of overlap. Very little or no overlap was found after injections of the premotor cortex (area 6), the visual areas 17, 18 and 19 and the auditory cortex (AI and AII).It is concluded that a major cortical input to the paramedian lobule arises in the posterior sigmoid gyrus (SmI), but that additional contributions arise in the anterior sigmoid gyrus (MsI), the parietal areas 5 and 7 and the second somatosensory cortex (SmII). Among the latter regions probably the parietal areas contribute most. Overlap between terminal regions of cortical fibres and cells projecting to the paramedian lobule takes place at numerous discrete sites at virtually all rostrocaudal levels of the pons. Cerebrocortical afferents via the pontine nuclei to the intermediate zone of the posterior lobe are organized according to the same principles as described previously for cortical afferents to the hemispheral parts of the posterior lobe (crus I and II).     

The organization of the projection from the cerebral cortex to the striatum in the rat

The detailed organization of the corticostriate projection has been investigated in the brain of the rat using the technique of retrograde transport of horseradish peroxidase following the placement of small, iontophoretic injections of horseradish peroxidase conjugated to lectin throughout all major regions of the striatum (caudate-putamen, nucleus accumbens and olfactory tubercle). The results demonstrate that all major regions of the cerebral cortex project to the striatum on both sides of the brain with an ipsilateral predominance. The cells of origin of both the ipsilateral and contralateral corticostriate projections lie mainly in lamina V (especially lamina Va) with very small numbers in lamina III of the neocortex and mesocortex, and in the deep laminae of the allocortex. The results show that each striatal locus receives inputs from several cortical regions, i.e. there is extensive overlap in the corticostriate projection, and that, in general terms, each cortical region projects onto a longitudinally oriented region of the striatum. In particular, the major subdivisions of the cerebral cortex--the neocortex, mesocortex and allocortex--project onto defined but partially overlapping regions of the striatum: the neocortex projects to the caudate-putamen; the mesocortex projects mainly to the medial and ventral regions of the caudate-putamen but also to the ventral striatum (nucleus accumens and olfactory tubercle); and the allocortex projects mainly to the ventral striatum but also to the medial and ventral parts of the caudate-putamen. Within each of these major projection systems there is a further organization, with the constituent parts of each major cortical region projecting to smaller longitudinal components of the major projection fields. Each neocortical area projects to a longitudinal region of the dorsal striatum (caudate-putamen): the sensory and motor areas project topographically onto the dorsolateral striatum such that the rostral sensorimotor cortex (head areas) projects to central and ventral regions and the more caudal sensorimotor cortex (limb areas) projects to dorsal regions of the dorsolateral striatum; the visual area projects to the dorsomedial striatum; and the auditory area projects to the medial striatum. Each mesocortical area projects to a longitudinal area of the striatum: the most posteromedial mesocortex (the retrosplenial area) projects to the dorsomedial striatum; more anterior and lateral parts of the mesocortex project to more ventral parts of the striatum: and the most lateral mesocortex (the agranular insular and perirhinal areas) project to the ventrolateral striatum.(ABSTRACT TRUNCATED AT 400 WORDS)     

Modular functional organization of cat anterior auditory field

Two tonotopic areas, the primary auditory cortex (AI) and the anterior auditory field (AAF), are the primary cortical fields in the cat auditory system. They receive largely independent, concurrent thalamocortical projections from the different thalamic divisions despite their hierarchical equivalency. The parallel streams of thalamic inputs to AAF and AI suggest that AAF neurons may differ from AI neurons in physiological properties. Although a modular functional organization in cat AI has been well documented, little is known about the internal organization of AAF beyond tonotopy. We studied how basic receptive field parameters (RFPs) are spatially organized in AAF with single- and multiunit recording techniques. A distorted tonotopicity with an underrepresentation in midfrequencies (1 and 5 kHz) and an overrepresentation in the high-frequency range was found. Spectral bandwidth (Q-values) and response threshold were significantly correlated with characteristic frequency (CF). To understand whether AAF has a modular organization of RFPs, CF dependencies were eliminated by a nonparametric, local regression model, and the residuals (difference between the model and observed values) were evaluated. In a given isofrequency domain, clusters of low or high residual RFP values were interleaved for threshold, spectral bandwidth, and latency, suggesting a modular organization. However, RFP modules in AAF were not expressed as robustly as in AI. A comparison of RFPs between AAF and AI shows that AAF neurons were more broadly tuned and had shorter latencies than AI neurons. These physiological field differences are consistent with anatomical evidence of largely independent, concurrent thalamocortical projections in AI and AAF, which strongly suggest field-specific processing.     

Frequency organization and responses to complex sounds in the medial geniculate body of the mustached bat

The auditory cortex of the mustached bat (Pteronotus parnellii) displays some of the most highly developed physiological and organizational features described in mammalian auditory cortex. This study examines response properties and organization in the medial geniculate body (MGB) that may contribute to these features of auditory cortex. About 25% of 427 auditory responses had simple frequency tuning with single excitatory tuning curves. The remainder displayed more complex frequency tuning using two-tone or noise stimuli. Most of these were combination-sensitive, responsive to combinations of different frequency bands within sonar or social vocalizations. They included FM-FM neurons, responsive to different harmonic elements of the frequency modulated (FM) sweep in the sonar signal, and H1-CF neurons, responsive to combinations of the bat's first sonar harmonic (H1) and a higher harmonic of the constant frequency (CF) sonar signal. Most combination-sensitive neurons (86%) showed facilitatory interactions. Neurons tuned to frequencies outside the biosonar range also displayed combination-sensitive responses, perhaps related to analyses of social vocalizations. Complex spectral responses were distributed throughout dorsal and ventral divisions of the MGB, forming a major feature of this bat's analysis of complex sounds. The auditory sector of the thalamic reticular nucleus also was dominated by complex spectral responses to sounds. The ventral division was organized tonotopically, based on best frequencies of singly tuned neurons and higher best frequencies of combination-sensitive neurons. Best frequencies were lowest ventrolaterally, increasing dorsally and then ventromedially. However, representations of frequencies associated with higher harmonics of the FM sonar signal were reduced greatly. Frequency organization in the dorsal division was not tonotopic; within the middle one-third of MGB, combination-sensitive responses to second and third harmonic CF sonar signals (60-63 and 90-94 kHz) occurred in adjacent regions. In the rostral one-third, combination-sensitive responses to second, third, and fourth harmonic FM frequency bands predominated. These FM-FM neurons, thought to be selective for delay between an emitted pulse and echo, showed some organization of delay selectivity. The organization of frequency sensitivity in the MGB suggests a major rewiring of the output of the central nucleus of the inferior colliculus, by which collicular neurons tuned to the bat's FM sonar signals mostly project to the dorsal, not the ventral, division. Because physiological differences between collicular and MGB neurons are minor, a major role of the tecto-thalamic projection in the mustached bat may be the reorganization of responses to provide for cortical representations of sonar target features.     

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.     

Afferent connections of dorsal and ventral agranular insular cortex in the hamsterMesocricetus auratus

The agranular insular cortex is transitional in location and structure between the ventrally adjacent olfactory allocortex primutivus and dorsally adjacent sensory-motor isocortex. Its ventral anterior division receives major afferent projections from olfactory areas of the limbic system (posterior primary olfactory cortex, posterolateral cortical amygdaloid nucleus and lateral entorhinal cortex) while its dorsal anterior division does so from non-olfactory limbic areas (lateral and basolateral amygdaloid nuclei).The medial segment of the mediodorsal thalamic nucleus projects to both the ventral and dorsal divisions of the agranular insular cortex, to the former from its anterior portion and to the latter from its posterior portion. Other thalamic inputs to the two divisions arise from the gelatinosus, central medial, rhomboid and parafascicular nuclei. The dorsal division, but not the ventral division, receives input from neurons in the lateral hypothalamus and posterior hypothalamus.The medial frontal cortex projects topographically and bilaterally upon both ventral and dorsal anterior insular cortex, to the former from the ventrally located medial orbital and infralimbic areas, to the latter from the dorsally-located anterior cingulate and medial precentral areas, and to both from the intermediately located prelimbic area. Similarly, the ipsilateral posterior agranular insular cortex and perirhinal cortex project in a topographic manner upon the two divisions of the agranular insular cortex.Commissural input to both divisions originates from pyramidal neurons in the respective contralateral homotopical cortical area. In each case, pyramidal neurons in layer V contribute 90% of this projection and 10% arises from layer III pyramidals.In the brainstem, the dorsal raphe nucleus projects to the ventral and dorsal divisions of the agranular insular cortex and the parabrachial nucleus projects to the dorsal division.Based on their cytoarchitecture, pattern of afferent connections and known functional properties, we consider the ventral and dorsal divisions of the agranular insular cortex to be, respectively, periallocortical and proisocortical portions of the limbic cortex.     

Microtopography of the dual corticothalamic projections originating from domains along the frequency axis of the cat primary auditory cortex

Spatial relationships between clusters of corticothalamic (CT) large terminals originating from cortical domains tuned to different frequencies were examined by pair-injecting two different anterograde tracers. Large-terminal CT projection originating from layer 5 was highly divergent with each injection site producing, on average, 15 local clusters distributing throughout non-lemniscal thalamic nuclei following a single anterograde tracer injection in the cat primary auditory cortex. Paired injections in higher- and lower-frequency cortical domains, resulting in labeling of two independent sets of terminal clusters, showed five recognizable patterns of spatial interaction between them. (1) In the ventral division of the medial geniculate complex (vMGC), sheet-like plexuses of small terminals of different origins were situated in parallel, with minimal overlap. (2) Extensive overlap of two low-density plexuses of differently labeled small terminals was observed in the medial division of the medial geniculate complex (MGC). (3) At the transition zones between the vMGC and the superficial dorsal nucleus of the MGC dorsal division, and between the vMGC and the ventrolateral nucleus, there were relatively broad clusters of a high density of large-terminal structures from the two cortical domains, which overlapped extensively. (4) At multiple loci in the nonlemniscal nuclei, pairing of two small clusters of differently labeled large terminals was observed. (5) Small unpaired clusters of large terminals were also found in the nonlemniscal nuclei. For large terminals, approximately 14%, 59%, and 27% clusters per injection demonstrated patterns 3, 4, and 5, respectively. The results provide evidence for the precise topographical organization for the large-terminal CT system at the microscopic level despite its highly divergent projection. This microtopographical projection from the tonotopic cortical field to non-tonotopic thalamic nuclei may raise the possibility of presence of a map that has not been defined in auditory non-lemniscal thalamic nuclei yet.     

Retinofugal projections of an echolocating megachiropteran

The projection leading from the eye and the nuclear targets of the projection to the brainstem were identified in an echolocating megachiropteran (Rousettus aegyptiacus) following unilateral intraocular injections of radioactive amino acids. In the hypothalamus, the projection ended bilaterally in suprachiasmatic nuclei. In the ventral thalamus, it ended bilaterally in external and internal divisions of the ventral lateral geniculate nuclei. In the dorsal thalamus, the projection terminated bilaterally in the dorsal lateral geniculate nuclei and contralaterally in the lateral posterior nucleus. Input from the two eyes was segregated to laminae in the lateral division of the dorsal lateral geniculate nucleus. The contralateral projection ended in the dorsolateral and ventral portions of lamina 1, in lamina 2, the ventral portions of lamina 3, and an interlaminar fiber plexus. The ipsilateral projection ended in the dorsomedial portion of lamina 1, the dorsal portion of lamina 3, and the most superficial portion of lamina 1. Contralateral and ipsilateral input to the medial division of the dorsal lateral geniculate nucleus was for the most part segregated. The projection to the pretectum terminated in nuclei of the optic tract, pretectal olivary nuclei, and posterior pretectal nuclei. Although the input to the pretectal nuclei was bilateral, the contralateral projection was greater. The contralateral projection to the superior colliculus terminated throughout the rostral-caudal extent of the superficial gray layer. The ipsilateral projection to the superior colliculus ended in the superficial gray layer in the middle one-third of the superior colliculus only. On the contralateral side the projection to the outer portion of the superficial gray layer was especially heavy. The superior fascicle of the accessory optic tract was identified. It was traced to dorsal, lateral, and medial accessory optic nuclei. These results indicate that the visual system of Rousettus is more extensive than that of the echolocating microchiroptera and that it is similar to that described for nonecholocating Pteropus.