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     

Branched projections in the auditory thalamocortical and corticocortical systems

Branched axons (BAs) projecting to different areas of the brain can create multiple feature-specific maps or synchronize processing in remote targets. We examined the organization of BAs in the cat auditory forebrain using two sensitive retrograde tracers. In one set of experiments (n=4), the tracers were injected into different frequency-matched loci in the primary auditory area (AI) and the anterior auditory field (AAF). In the other set (n=4), we injected primary, non-primary, or limbic cortical areas. After mapped injections, percentages of double-labeled cells (PDLs) in the medial geniculate body (MGB) ranged from 1.4% (ventral division) to 2.8% (rostral pole). In both ipsilateral and contralateral areas AI and AAF, the average PDLs were <1%. In the unmapped cases, the MGB PDLs ranged from 0.6% (ventral division) after insular cortex injections to 6.7% (dorsal division) after temporal cortex injections. Cortical PDLs ranged from 0.1% (ipsilateral AI injections) to 3.7% in the second auditory cortical area (AII) (contralateral AII injections). PDLs within the smaller (minority) projection population were significantly higher than those in the overall population. About 2% of auditory forebrain projection cells have BAs and such cells are organized differently than those in the subcortical auditory system, where BAs can be far more numerous. Forebrain branched projections follow different organizational rules than their unbranched counterparts. Finally, the relatively larger proportion of visual and somatic sensory forebrain BAs suggests modality specific rules for BA organization.     

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.     

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.     

Functional organization of the callosal connections of the cat auditory cortex

In acute experiments on immobilized cats, using a method of topographical recording of homotopic and heterotopic transcallosal responses, the functional organization of the callosal connections of the auditory cortex was investigated. It was established that the homotopic potentials of the primary projection field (AI) have the greatest amplitude, minimal temporal parameters, and the maximal stability of these characteristics as compared with the associative fields of the auditory cortex (AII, AIV, Ep). The heterotropic transcallosal responses in field AI appeared during stimulation of the analogous field, while in field Ep, they were recorded both during stimulation of the analogous field, and of fields AI and AII of the opposite hemisphere. It is hypothesized that the structure of the transcallosal connections of the primary projection fields of the auditory cortex is characterizised by homotopy, whereas in the associative auditory fields the role of heterotopic transcallosal interactions increases. It is possible that such a structure of the transcallosal connections assures a significant role for interhemispheric interactions in the mechanisms of spatial audition.Translated from Fiziologicheskii Zhurnal SSSR imeni I. M. Sechenova, Vol. 73, No. 7, pp. 860–867, July, 1987.     

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.     

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.     

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   

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.     

Efferent connections of "posterodorsal" auditory area in the rat cortex: implications for auditory spatial processing

We examined efferent connections of the cortical auditory field that receives thalamic afferents specifically from the suprageniculate nucleus (SG) and the dorsal division (MGD) of the medial geniculate body (MG) in the rat [Neuroscience 117 (2003) 1003]. The examined cortical region was adjacent to the caudodorsal border (4.8-7.0 mm posterior to bregma) of the primary auditory area (area Te1) and exhibited relatively late auditory response and high best frequency, compared with the caudal end of area Te1. On the basis of the location and auditory response property, the cortical region is considered identical to "posterodorsal" auditory area (PD). Injections of biocytin in PD revealed characteristic projections, which terminated in cortical areas and subcortical structures that play pivotal roles in directed attention and space processing. The most noticeable cortical terminal field appeared as dense plexuses of axons in area Oc2M, the posterior parietal cortex. Small terminal fields were scattered in area frontal cortex, area 2 that comprises the frontal eye field. The subcortical terminal fields were observed in the pontine nucleus, the nucleus of the brachium inferior colliculus, and the intermediate and deep layers of the superior colliculus. Corticostriatal projections targeted two discrete regions of the caudate putamen: the top of the middle part and the caudal end. It is noteworthy that the inferior colliculus and amygdala virtually received no projection. Corticothalamic projections terminated in the MGD, the SG, the ventral zone of the ventral division of the MG, the ventral margin of the lateral posterior nucleus (LP), and the caudodorsal part of the posterior thalamic nuclear group (Po). Large terminals were found in the MGD, SG, LP and Po besides small terminals, the major component of labeling. The results suggest that PD is an auditory area that plays an important role in spatial processing linked to directed attention and motor function. The results extend to the rat findings from nonhuman primates suggesting the existence of a posterodorsal processing stream for auditory spatial perception.     

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.     

Auditory thalamic nuclei projections to the temporal cortex in the rat

Thalamocortical projections from the auditory thalamic nuclei were examined systematically in the rat, including those from the dorsal division (MGD) of the medial geniculate body (MG), which were less clearly determined in previous studies. Injections of biocytin confined in each thalamic nucleus revealed characteristic features of projections in terms of cortical areas and layers of termination. In contrast to exclusively selective projections to cortical area Te1 from the ventral division (MGV) of the MG, diffuse and selective terminations were observed in the projections from the dorsal (MGD) and medial divisions (MGM) of the MG and the suprageniculate nucleus (SG). Diffuse termination was continuous in layer I or VI of the temporal cortex, while selective termination was in layers III and IV of discrete cortical areas. In addition to diffuse termination in the upper half of layer I of cortical areas Te1, Te2d and Te3v, the MGD and SG projections formed plexuses of axons selectively in lower layer III and layer IV of Te2d and Te3v. The SG projections targeted further the dorsal bank of the perirhinal cortex (PRh), while the MGD projections targeted in part the ventral fringe of Te1. The MGM projections terminated diffusely in layer VI of Te1 and Te3v, and selectively in lower layer III and layer IV of the rostral part of Te3v. Diffuse projections to layers I and VI from the SG and MGM extended in cortical regions over the dorsal fringe of Te1. Selective dense projections to middle cortical layers of Te2d and Te3v (especially its rostral part) indicate the existence of auditory areas, which could be involved in cross-modal interaction with visual and somatosensory system, respectively. Diffuse projections are supposed to bind information processings in these areas and the primary auditory area (Te1).     

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.     

Inputs from the olfactory bulb and olfactory cortex to the entorhinal cortex in the cat

Summary The spatial organization and laminar distribution of projections from the olfactory bulb and the anterior (PPCa) and posterior (PPCp) divisions of the prepiriform cortex to the entorhinal cortex were studied with anterograde (³H-leucine) and retrograde (WGA-HRP) tracing techniques. After ³H-leucine injections into the olfactory bulb transported labeling was seen over the lateral entorhinal area, except its most medial part, and over the rostral part of the medial entorhinal area. The labeling covers exclusively layer Ia. The lateral and medial entorhinal areas are also reached by fibers from the prepiriform cortex. The projection to the medial entorhinal area has not been described previously. Following injections of ³H-leucine into the PPCa transported labeling is present over the entire expanse of the entorhinal cortex and is located over layer Ib with the greatest density in its superficial part. Injections of ³H-leucine into the PPCp give rise to transported labeling over much of the entorhinal cortex. No labeling was found over the most medial parts of the medial subdivision (VMEA) of the lateral entorhinal area and the medial entorhinal area. Labeling occupies layer Ib, especially its middle part, and layers II and III. Both PPCa and PPCp appear to project most heavily to the dorsal (DLEA) and ventral (VLEA) subdivisions of the lateral entorhinal area. From the retrograde experiments it can be inferred that cells of layers II and III of the PPCa project predominantly to the DLEA, whereas those of the PPCp project predominantly to the VLEA. The MEA receives its heaviest projection from layer II of both PPCa and PPCp. In control experiments with ³H-leucine injections into the endopiriform nucleus it was found that this nucleus projects to the entire expanse of the entorhinal cortex. The fibers distribute to all layers with the exception of layer Ia.Abbreviations AI agranular insular cortex - AL lateral nucleus of the amygdala - BL basolateral nucleus of the amygdala - BM basomedial nucleus of the amygdala - C claustrum - CoA cortical nucleus of the amygdala - DLEA dorsal division of the lateral entorhinal cortex - END endopiriform nucleus - H hippocampus - I granular insular cortex - lot lateral olfactory tractus - MCL mitral cell layer of the olfactory bulb - MEA medial entorhinal area - OB olfactory bulb - PPCa anterior part of the prepiriform nucleus - PPCp posterior part of the prepiriform nucleus - VLEA ventral division of the lateral entorhinal cortex - VMEA ventromedial division of the lateral entorhinal cortex - 35 area 35 of the perirhinal cortex - 36 area 36 of the perirhinal cortex     

Topography of projections from the primary and non-primary auditory cortical areas to the medial geniculate body and thalamic reticular nucleus in the rat

The functional significance of parallel and redundant information processing by multiple cortical auditory fields remains elusive. A possible function is that they may exert distinct corticofugal modulations on thalamic information processing through their parallel connections with the medial geniculate body and thalamic reticular nucleus. To reveal the anatomical framework for this function, we examined corticothalamic projections of tonotopically comparable subfields in the primary and non-primary areas in the rat auditory cortex. Biocytin was injected in and around cortical area Te1 after determining best frequency at the injection site on the basis of epicortical field potentials evoked by pure tones. The rostral part of area Te1 (primary auditory area) and area temporal cortex, area 2, dorsal (Te2D) (posterodorsal auditory area) dorsal to the caudal end of area Te1, which both exhibited high best frequencies, projected to the ventral zone of the ventral division of the medial geniculate body. The caudal end of area Te1 (auditory area) and the rostroventral part of area Te1 (a part of anterior auditory field), which both exhibited low best frequencies, projected to the dorsal zone of the ventral division of the medial geniculate body. In contrast to the similar topography in the projections to the ventral division of the medial geniculate body, collateral projections to the thalamic reticular nucleus terminated in the opposite dorsal and ventral zones of the lateral and middle tiers of the nucleus in each pair of the tonotopically comparable cortical subfields. In addition, the projections of the non-primary cortical subfields further arborized in the medial tier of the thalamic reticular nucleus. The results suggest that tonotopically comparable primary and non-primary subfields in the auditory cortex provide corticofugal excitatory effects to the same part of the ventral division of the medial geniculate body. On the other hand, corticofugal inhibition via the thalamic reticular nucleus may operate in different parts of the ventral division of the medial geniculate body or different thalamic nuclei. The primary and non-primary cortical auditory areas are presumed to subserve distinct gating functions for auditory attention.     

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.     

Auditory cortical field projections to the basal ganglia of the cat

Projections to the basal ganglia from four auditory cortical fields in the cat were studied by combining microelectrode-mapping of the neurons' best frequencies with autoradiographic and histochemical tract-tracing techniques. Each auditory field is a source of projections to the homolateral basal ganglia. The distribution of labeling within the basal ganglia is related to the cortical field in which the injection site is located. The dorsal portion of the putamen and adjacent caudate nucleus are connected with cortical fields situated anteriorly and dorsally, while the ventral portion of the putamen and adjacent lateral amygdaloid nucleus are related to auditory fields situated posteriorly and ventrally. Injections of two different tracers into different best-frequency loci of one cortical field provided evidence that low best-frequency neurons project medially within the basal ganglia while high best-frequency neurons project more laterally.We concluded that there was a basic similarity among patterns of terminations in the basal ganglia from axons that originate in different auditory cortical fields. When the source of a projection was confined to a restricted portion of an auditory cortical field, labeling appeared as dense patches of silver grains separated from each other by areas of less dense labeling. Often, these patches were distributed within a sheet of tissue, elongated both dorsoventrally and anteroposteriorly. Loci having the same best-frequency representation, but situated in different auditory cortical fields, project upon overlapping but not coextensive portions of a single sheet of tissue. Thus the projections from geographically distant cortical loci possessing similar best-frequency representations are notably distinguished on a topographic basis. By comparison, two adjacent sheets of tissue were labeled when two injections were made into the low best-frequency and high best-frequency representations of the same auditory field. Doubleinjection, double-tracer experiments revealed that adjacent sheets of tissue received projections from different best-frequency loci. These observations suggested a degree of tonotopic organization to this projection system which was equipoise to the evidence obtained for a topographic organization.     

Restricted cortical termination fields of the midline and intralaminar thalamic nuclei in the rat.

The projections from the midline and intralaminar thalamic nuclei to the cerebral cortex were studied in the rat by means of anterograde tracing with Phaseolus vulgaris-leucoagglutinin. The midline and intralaminar nuclear complex taken as a whole projects to widespread, predominantly frontal, cortical areas. Each of the constituent thalamic nuclei has a restricted cortical projection field that overlaps only slightly with the projection fields of adjacent midline and intralaminar nuclei. The projections of the intralaminar nuclei cover a larger cortical area than those of the midline nuclei. The laminar distributions of fibres from individual midline and intralaminar thalamic nuclei are different and include both deep and superficial cortical layers. The parataenial, paraventricular and intermediodorsal midline nuclei each project to circumscribed parts of the prefrontal cortex and the hippocampal and parahippocampal regions. In the prefrontal cortex, the projections are restricted to the medial orbital, infralimbic, ventral prelimbic and agranular insular fields, and the rostral part of the ventral anterior cingular cortex. In contrast to the other midline nuclei, the rhomboid nucleus projects to widespread cortical areas. The rostral intralaminar nuclei innervate dorsal parts of the prefrontal cortex, i.e. the dorsal parts of the prelimbic, anterior cingular and dorsal agranular insular cortical fields, the lateral and ventrolateral orbital areas, and the caudal part of the ventral anterior cingular cortex. Additional projections are aimed at the agranular fields of the motor cortex and the caudal part of the parietal cortex. The lateral part of the parafascicular nucleus sends fibres predominantly to the lateral agranular field of the motor cortex and the rostral part of the parietal cortex. The medial part of the parafascicular nucleus projects rather sparsely to the dorsal part of the prelimbic cortex, the anterior cingular cortex and the medial agranular field of the motor cortex. Individual midline and intralaminar thalamic nuclei are thus in a position to directly influence circumscribed areas of the cerebral cortex. In combination with previously reported data on the organization of the midline and intralaminar thalamostriatal projections and the prefrontal corticostriatal projections the present results suggest a high degree of differentiation in the convergence of thalamic and cortical afferent fibres in the striatum. Each of the recently described parallel basal ganglia-thalamocortical circuits can thus be expanded to include projections at both the cortical and striatal levels from a specific part of the midline and intralaminar nuclear complex. The distinctive laminar distributions of the fibres originating from the different nuclei emphasize the specificity of the midline and intralaminar thalamocortical projections.     

Extracortical descending projections to the rat inferior colliculus

Feedback controlling is an important element in the sensory processing in the auditory system. It has been long recognized that the inferior colliculus (IC) sends direct ascending projections to the medial geniculate body (MGB), but receives feedback regulation from the auditory cortex. In the present study we probed the shorter extracortical projections to the IC, including the direct descending pathway from the MGB. In the rat, the fluorescence retrograde tracers Fluorogold, True Blue or Rhodamine latex microspheres were injected into the IC, and the auditory thalamus and surrounding regions were examined for fluorescent neurones. We did not find any retrograde labelling in the ventral division of the MGB. However, retrogradely labelled neurones were found in the medial and suprageniculate nuclei of the MGB. We also observed densely packed groups of fluorescent neurones in the peripeduncular nucleus and numerous labelled neurones in the nucleus of the brachium of the IC. The existence of a direct descending pathway to the IC from at least some auditory thalamic nuclei challenges the perception of the colliculo-thalamic relationship as one-way traffic and suggests more direct involvement of the auditory thalamus in the feedback regulation of the incoming acoustic signals.