Anatomy of the auditory thalamocortical system of the guinea pig

We investigated the projection from the medial geniculate body (MG) to the tonotopic fields (the anterior field A, the dorsocaudal field DC, the small field S) and to the nontonotopic ventrocaudal belt in the auditory cortex of the guinea pig. The auditory fields were first delimited in electrophysiological experiments with microelectrode mapping techniques. Then, small quantities of horseradish peroxidase (HRP) and/or fluorescent retrograde tracers were injected into the sites of interest, and the thalamus was checked for labeled cells. The anterior field A receives its main thalamic input from the ventral nucleus of the MG (MGv). The projection is topographically organized. Roughly, the caudal part of the MGv innervates the rostral part of field A and vice versa. After injection of tracer into low or medium best-frequency sites in A, we also found a topographic gradient along the isofrequency contours: the dorsal (ventral) part of a cortical isofrequency strip receives afferents from the rostral (caudal) portions of the corresponding thalamic isofrequency band. However, it is not so obvious whether such a gradient exists also in the high-frequency part of the projection. A second, weaker projection to field A originates in a magnocellular nucleus that is situated caudomedially in the MG and was therefore named the caudomedial nucleus. The dorsocaudal field DC receives input from the same nuclei as the anterior field, but the location of the labeled cells in the MGv is different. This was demonstrated by injection of different tracers into sites with like best frequencies in fields A and DC, respectively. After injection of HRP into the 1-2-kHz isofrequency strip in field A and injection of Nuclear Yellow (NY) into the 1-2-kHz site in field DC, the labeled cells in the MGv form one continuous array that runs from caudal to rostral over the whole extent of the MGv. The anterior part of this array consists of NY-labeled cells; i.e., it projects to field DC. The caudal part is formed by HRP-labeled cells; i.e., it innervates field A. These findings indicate that there is only one continuous tonotopic map in the MGv. This map is split when projected onto the cortex so that two adjacent tonotopic fields (A and DC) result. The cortical maps are rotated relative to the thalamic map in that rostral portions of the MGv project to caudal parts of the tonotopic cortex and vice versa.(ABSTRACT TRUNCATED AT 400 WORDS)     

Thalamic connectivity of the second somatosensory area and neighboring somatosensory fields of the lateral sulcus of the macaque

The thalamocortical relations of the somatic fields in and around the lateral sulcus of the macaque were studied following cortical injections of tritated amino acids and horseradish peroxidase (HRP). Special attention was paid to the second somatosensory area (S2), the connections of which were also studied by means of thalamic isotope injections and retrograde degeneration. S2 was shown to receive its major thalamic input from the ventroposterior inferior thalamic nucleus (VPI) and not, as previously reported, from the caudal division of the ventroposterior lateral nucleus (VPLc). Following small injections of isotope or HRP into the hand representation of S2, only VPI was labeled. Larger injections, which included the representations of more body parts, led to heavy label in VPI, with scattered label in VPLc, the central lateral nucleus (CL), and the posterior nucleus (Po). In addition, small isotope injections into VPLc did not result in label in S2 unless VPI was also involved in the injection site, and ablations of S2 led to cell loss in VPI. Comparison of injections involving different body parts in S2 suggested a somatotopic arrangement within VPI such that the trunk and lower limb representations are located posterolaterally and the hand and arm representations anteromedially. The location of the thalamic representations of the head, face, and intraoral structures that project to S2 may be in the ventroposterior medial nucleus (VPM). The granular (Ig) and dysgranular (Id) fields of the insula and the retroinsular field (Ri) each receive inputs from a variety of nuclei located at the posteroventral border of the thalamus. Ig receives its heaviest input from the suprageniculate-limitans complex (SG-Li), with additional inputs from Po, the magnocellular division of the medial geniculate n. (MGmc), VPI, and the medial pulvinar (Pulm). Id receives its heaviest input from the basal ventromedial n. (VMb), with additional inputs from VPI, Po, SG-Li, MGmc, and Pulm. Ri receives its heaviest input from Po, with additional input from SG-Li, MGmc, Pulm, and perhaps VPI. Area 7b receives its input from Pulm, the oral division of the pulvinar, the lateral posterior n., the medial dorsal n., and the caudal division of the ventrolateral n. These results indicate that the somatic cortical fields, except for those comprising the first somatosensory area, each receive inputs from an array of thalamic nuclei, rather than just one, and that individual thalamic somatosensory relay nuclei each project to more than one cortical field.     

Visual and auditory association areas of the cat's posterior ectosylvian gyrus: thalamic afferents

The feline posterior ectosylvian gyrus contains a broad band of association cortex that is bounded anteriorly by tonotopic auditory areas and posteriorly by retinotopic visual areas. To characterize the possible functions of this cortex and to throw light on its pattern of internal divisions, we have carried out an analysis of its thalamic afferents. Deposits of differentiable retrograde tracers were placed at 17 cortical sites in nine cats. The deposit sites spanned the crown of the posterior ectosylvian gyrus and adjacent cortex in the suprasylvian sulcus. We compiled counts of retrogradely labeled neurons in 12 thalamic nuclei delineated by use of Nissl and acetylcholinesterase stains. We then employed a statistical clustering algorithm to identify groups of injections that gave rise to similar patterns of thalamic labeling. The results suggest that the posterior ectosylvian gyrus contains 3 fundamentally different cortical districts that have the form of parallel vertical bands. Very anterior cortex, overlapping previously identified tonotopic auditory areas (AI, P and VP) receives a dense projection from the laminated division of the medial geniculate body (MGl). An intermediate strip, to which we refer as the auditory belt, is innervated by axons from nontonotopic divisions of the medial geniculate body (MGds, MGvl, MGm, and MGd), from the lateral division of the posterior group (Pol), and from the posterior suprageniculate nucleus (SGp). A posterior strip, to which we refer as EPp, receives strong projections from the LM-SG complex (LM-SGa and LMp), and lighter projections from the intralaminar and lateroposterior (LPm and LPl) nuclei. On grounds of thalamic connectivity, EPp is not obviously distinguishable from adjacent retinotopic visual areas (PLLS, DLS, and VLS), and may be regarded as forming, together with these areas, a connectionally homogeneous visual belt.     

Visual and auditory association areas of the cat's posterior ectosylvian gyrus: cortical afferents

In a preceding report, we described patterns of thalamic retrograde labeling following 17 tracer deposits on the cat's posterior ectosylvian gyrus and concluded, on the basis of patterns of thalamic connectivity, that the posterior ectosylvian gyrus is composed of three major divisions: a tonotopic auditory zone located anteriorly, a belt of auditory association cortex occupying the gyral crown, and a visual belt located posteriorly. We describe here patterns of transcortical retrograde labeling obtained from tracer deposits in the three zones so defined. Our results indicate that the tonotopic auditory strip is innervated primarily by axons from low-order auditory areas (AAF, AI, P, VP, and V), that the auditory belt receives its strongest input from nontonotopic auditory fields (AII, temporal cortex, and other parts of the auditory belt), and that projections to the visual belt derive primarily from extrastriate visual areas (ALLS, PLLS, DLS, 19, 20, and 21) and from association areas affiliated with the visual system (insular cortex, posterior cingulate gyrus, area 7p, and frontal cortex). We discuss the results in relation to previous systems for parcellating the posterior ectosylvian gyrus of the cat and consider the possibility that divisions of the feline posterior ectosylvian gyrus correspond directly to areas making up the superior temporal gyrus in primates.     

Organization of thalamic projections to the ventral striatum in the primate

Although thalamic projections to the dorsal striatum are well described in primates and other species, little is known about thalamic projections to the ventral or “limbic” striatum in the primate. This study explores the organization of the thalamic projections to the ventral striatum in the primate brain by means of wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP) and Lucifer yellow (LY) retrograde tracer techniques. In addition, because functional and connective differences have been described for the core and shell components of the nucleus accumbens in the rat and are thought to be similar in the primate, this study also explores whether these regions of the nucleus accumbens can be distinguished by their thalamic input. Tracer injections are placed in different portions of the ventral striatum, including the medial and lateral regions of the ventral striatum; the central region of the ventral striatum, including the dorsal part of the core of the nucleus accumbens; and the shell region of the nucleus accumbens. Retrogradely labeled neurons are located mainly in the midline nuclear group (anterior and posterior paraventricular, paratenial, rhomboid, and reuniens thalamic nuclei) and in the parafascicular thalamic nucleus. Additional labeled cells are found in other portions of the intralaminar nuclear group as well as in other thalamic nuclei in the ventral, anterior, medial, lateral, and posterior thalamic nuclear groups. The distribution of labeled cells varies depending on the area of the ventral striatum injected. All regions of the ventral striatum receive strong projections from the midline thalamic nuclei and from the parafascicular nucleus. In addition, the medial region of the ventral striatum receives numerous projections from the central superior lateral nucleus, the magnocellular subdivision of the ventral anterior nucleus, and parts of the mediodorsal nucleus. After injection into the lateral region of the ventral striatum, few labeled neurons are seen scattered in nuclei of the intralaminar and ventral thalamic groups and occasional labeled cells in the mediodorsal nucleus. The central region of the ventral striatum, including the dorsal part of the core of the nucleus accumbens, receives a limited projection from the midline thqlamic, predominantly from the rhomboid nucleus. It receives much smaller projections from the central medial nucleus and the ventral, anterior, and medial thalamic groups. The shell of the nucleus accumbens receives the most limited projection from the thalamus and is innervated almost exclusively by the midline thalamic nuclei and the central medial and parafascicular nuclei. The shell is distinguished from the rest of the ventral striatum in that it receives the fewest projections from the ventral, anterior, medial, and lateral thalamic nuclei. © 1995 Wiley-Liss, Inc.     

Telencephalic connections in lizards. II. Projections to anterior dorsal ventricular ridge

Three distinct cytoarchitectonic regions were identified within the anterior dorsal ventricular ridge (ADVR) of two species of lizards, Gekko gecko and Iguana iguana. These regions have been named according to their general topographical positions: medial area, caudolateral area, and rostrolateral area. Injections of horseradish peroxidase throughout the ADVR demonstrated that each of the three areas of the ADVR receives projections from specific thalamic nuclei which are associated with specific sensory modalities. The medial area receives an auditory thalamic projection from nucleus medialis. The caudolateral area receives thalamic projections from nucleus medialis posterior and nucleus posterocentralis. The latter two nuclei were shown to receive projections from the spinal cord and, therefore, are presumed to be associated with body somatosensory information. The rostrolateral area receives a thalamic projection from nucleus rotundus, which receives visual information. In addition, the mesencephalic tegmentum and the thalamic nucleus dorsomedialis project to the entire ADVR. The latter projection is similar to the diffuse cortical projections of the intralaminar thalamic nuclei in mammals. These findings support previous suggestions that the ADVR is comparable to sensory regions of the mammalian neocortex.     

Organization of the thalamostriatal projections in the rat, with special emphasis on the ventral striatum

The organization of the thalamic projections to the ventral striatum in the rat was studied by placing injections of the retrograde tracer cholera toxin subunit B in the ventral striatum and small deposits of the anterograde tracer Phaseolus vulgaris-leucoagglutinin (PHA-L) in individual midline and intralaminar thalamic nuclei. In order to provide a complete map of the midline and intralaminar thalamostriatal projections, PHA-L injections were also made in those parts of the intralaminar nuclei that project to the dorsal striatum. The relationship of thalamic afferent fibres with the compartmental organization of the ventral striatum was assessed by combining PHA-L tracing and enkephalin immunohistochemistry. The various midline and intralaminar thalamic nuclei project to longitudinally oriented striatal sectors. The paraventricular thalamic nucleus sends most of its fibres to medial parts of the nucleus accumbens and the olfactory tubercle, whereas smaller contingents of fibres terminate in the lateral part of the nucleus accumbens and the most ventral, medial, and caudal parts of the caudate-putamen complex. The projections of the parataenial nucleus are directed towards central and ventral parts of the nucleus accumbens and intermediate mediolateral parts of the olfactory tubercle. The intermediodorsal nucleus projects to lateral parts of the nucleus accumbens and the olfactory tubercle and to ventral parts of the caudate-putamen. The projection of the rhomboid nucleus is restricted to the rostrolateral extreme of the striatum. A diffuse projection to the ventral striatum arises from neurons ventral and caudal to the nucleus reuniens rather than from cells inside the nucleus. Fibres from the central medial nucleus terminate centrally and dorsolaterally in the rostral part of the nucleus accumbens and medially in the caudate-putamen. Successively more lateral positions in the caudate-putamen are occupied by fibres from the paracentral and central lateral nuclei, respectively. The lateral part of the parafascicular nucleus projects to the most lateral part of the caudate-putamen, whereas projections from the medial part of this nucleus terminate in the medial part of the caudate-putamen and in the dorsolateral part of the nucleus accumbens. Furthermore, a rostral to caudal gradient in a midline or intralaminar nucleus corresponds to a dorsal to ventral and rostral to caudal gradient in the striatum. In the ventral striatum, thalamic afferent fibres in the "shell" region of the nucleus accumbens avoid areas of high cell density and weak enkephalin immunoreactivity.(ABSTRACT TRUNCATED AT 400 WORDS)     

Thalamic projections to retrosplenial cortex in the rat

The topographic relationships between anterior thalamic neurons and their terminal projection fields in the retrosplenial cortex of the rat were characterized by experiments with the fluorescent dye retrograde labeling technique. The results demonstrate that the anterodorsal (DAD) and anteroventral (AV) nuclei project heavily to retrosplenial granular cortex (Rg) and to a lesser extent to retrosplenial agranular cortex (Rag). In contrast, the anteromedial (AM) and lateral dorsal (LD) nuclei project heavily to Rag and more lightly to Rg. Irrespective of terminal field in Rg or Rag, the neuronal cell bodies in AD and AV are organized topographically so that the neurons in the caudal part of each nucleus project to rostral retrosplenial cortex and the neurons in the rostral portion of each nucleus project to the caudal retrosplenial cortex. Further, the ventromedial AD and AV neurons project to rostral retrosplenial cortex, whereas dorsolateral neurons in both nuclei project to caudal retrosplenial cortex. LD neurons display a different topographic organization. The neurons in the medioventral part of LD project primarily to the rostral retrosplenial cortex, and the neurons in lateral LD project to the caudal retrosplenial cortex. This latter projection to the caudal retrosplenial cortex is also contributed to by neurons residing in the mediodorsal part of caudal LD. The neurons in AM that project to the retrosplenial cortex display less segregation than the AV, AD, or LD neurons. In all experiments, a number of neurons in the dorsal ventro-anterolateral nucleus were labeled by retrosplenial injections. The largest number of cells in this nucleus were labeled after Rag injections, and these were topographically organized such that the neurons projecting to the rostral Rag were located immediately deep to the internal medullary lamina, and the neurons projecting to the caudal Rag were more ventrally located. Very few thalamic neurons have axon collaterals to different areas of the retrosplenial cortex as shown by double labeling experiments. Together, these results demonstrate a highly organized thalamic projection to the retrosplenial cortex.     

Organization of the Visual Reticular Thalamic Nucleus of the Rat

The visual sector of the reticular thalamic nucleus has come under some intense scrutiny over recent years, principally because of the key role that the nucleus plays in the processing of visual information. Despite this scrutiny, we know very little of how the connections between the reticular nucleus and the different areas of visual cortex and the different visual dorsal thalamic nuclei are organized. This study examines the patterns of reticular connections with the visual cortex and the dorsal thalamus in the rat, a species where the visual pathways have been well documented. Biotinylated dextran, an anterograde and retrograde tracer, was injected into different visual cortical areas [17; rostral 18a: presumed area AL (anterolateral); caudal 18a: presumed area LM (lateromedial); rostral 18b: presumed area AM (anteromedial); caudal 18b: presumed area PM (posteromedial)] and into the different visual dorsal thalamic nuclei (posterior thalamic, lateral posterior, lateral geniculate nuclei), and the patterns of anterograde and retrograde labelling in the reticular nucleus were examined. From the cortical injections, we find that the visual sector of the reticular nucleus is divided into subsectors that each receive an input from a distinct visual cortical area, with little or no overlap. Further, the resulting pattern of cortical terminations in the reticular nucleus reflects largely the patterns of termination in the dorsal thalamus. That is, each cortical area projects to a largely distinct subsector of the reticular nucleus, as it does to a largely distinct dorsal thalamic nucleus. As with each of the visual cortical areas, each of the visual dorsal thalamic (lateral geniculate, lateral posterior, posterior thalamic) nuclei relate to a separate territory of the reticular nucleus, with little or no overlap. Each of these dorsal thalamic territories within the reticular nucleus receives inputs from one or more of the visual cortical areas. For instance, the region of the reticular nucleus that is labelled after an injection into the lateral geniculate nucleus encompasses the reticular regions which receive afferents from cortical areas 17, rostral 18b and caudal 18b. These results suggest that individual cortical areas may influence the activity of different dorsal thalamic nuclei through their reticular connections.     

The subcortical auditory structures in the Mongolian gerbil: II. Frequency‐related topography of the connections with cortical field AI

We investigated the frequency‐related topography of connections of the primary auditory cortical field (AI) in the Mongolian gerbil with subcortical structures of the auditory system by means of the axonal transport of two bidirectional tracers, which were simultaneously injected into regions of AI with different best frequencies (BFs). We found topographic, most likely frequency‐matched (tonotopic) connections as well as non‐topographic (non‐tonotopic) connections. AI projects in a tonotopic way to the ipsilateral ventral (MGv) and dorsal divisions (MGd) of the medial geniculate body (MGB), the reticular thalamic nucleus and dorsal nucleus of the lateral lemniscus, and the ipsi‐ and contralateral dorsal cortex of the inferior colliculus (IC) and central nucleus of the IC. AI receives tonotopic inputs from MGv and MGd. Projections from different BF regions of AI terminate in a non‐tonotopic way in the ipsilateral medial division of the MGB (MGm), the suprageniculate thalamic nucleus (SG) and brachium of the IC (bic), and the ipsi‐ and contralateral external cortex and pericollicular areas of the IC. The anterograde labeling in the intermediate and ventral nucleus of the lateral lemniscus, parts of the superior olivary complex, and divisions of the cochlear nucleus was generally sparse; thus a clear topographic arrangement of the labeled axons could not be ruled out. AI receives non‐tonotopic inputs from the ipsilateral MGm, SG, and bic. In conclusion, the tonotopic and non‐tonotopic corticofugal connections of AI can potentially serve for both conservation and integration of frequency‐specific information in the respective target structures. J. Comp. Neurol. 521:2772–2797, 2013. © 2013 Wiley Periodicals, Inc.     

Descending projections from the auditory cortex to the inferior colliculus in the gerbil, Meriones unguiculatus

Corticofugal projections to the auditory midbrain, the inferior colliculus (IC), influence the way in which specific sets of IC neurons process acoustic signals. We used retrograde tracer (Fluorogold, Fluororuby, microbeads) injections in the IC to study the morphology and location of cortico-collicular projecting neurons and anterograde tracer (dextran biotin) injections in auditory cortical fields to describe the distribution of terminals in the IC. Nissl staining, cytochrome oxidase activity, and neurofilament SMI32 immunostaining were used to delimit the different auditory areas. We defined a primary or "core" auditory cortex and a secondary "caudal" auditory area containing layer V pyramidal neurons that project to the IC. These projections target the central nucleus of the IC (CNIC) ipsilaterally and the IC cortices bilaterally, with the ipsilateral component predominant. Other secondary auditory areas, dorsal and ventral to the core, do not directly participate in this projection. The ventral secondary cortex targets midbrain periaqueductal gray. The projection from the core cortex originates from two classes of layer V pyramidal cells. Cells presenting a tufted apical dendrite in layer I have dense terminal fields in the IC cortices. Pyramids lacking layer I dendritic tufts target the CNIC in a less dense but tonotopic manner. The caudal cortex projection originates from smaller layer V pyramids and targets the IC cortices with dense terminal fields. Descending auditory inputs from the core and caudal areas converge in the dorsal and external cortices of the IC. Descending connections to the gerbil IC form a segregated system in which multiple descending channels originating from different neuronal subpopulations may modulate specific aspects of ascending auditory information.     

Auditory cortical projections to the cat inferior colliculus

The projection from 11 auditory cortical areas onto the subdivisions of the inferior colliculus was studied in adult cats by using two different anterograde tracers to label corticocollicular (CC) axon terminals. The main results were that: 1) a significant CC projection arose from every field; 2) the principal inferior collicular targets were the dorsal cortex, lateral nucleus, caudal cortex, and intercollicular tegmentum, with only a sparse projection to the central nucleus; 3) the input was usually bilateral, with the ipsilateral side by far the most heavily labeled, and the contralateral projection was a symmetrical subset of the ipsilateral input; 4) the CC system is both divergent and convergent, with single cortical areas projecting to six or more collicular subdivisions, and each auditory midbrain subdivision receiving a convergent projection from two to ten cortical areas; 5) cortical areas devoid of tonotopic organization have topographic projections to collicular target nuclei; 6) the heaviest CC projection terminated in the caudal half of the inferior colliculus; and finally, 7) the relative strength of the corticocollicular labeling was far less than that of the corresponding corticothalamic projection in the same experiments. The CC system is strategically placed to influence both descending and ascending pathways arising in the inferior colliculus. Nuclei that participate in the premotor system, like the inferior collicular subdivisions that project to the pons, receive substantial corticofugal input. Both the dorsal (pericentral) and the lateral (external) nuclei of the inferior colliculus project to parts of the medial geniculate body whose closest auditory affiliations are with nontonotopic cortical regions involved in higher order auditory perception. The corticocollicular system may link brainstem and colliculothalamic circuits to coordinate premotor and perceptual aspects of hearing. J. Comp. Neurol. 400:147–174, 1998. © 1998 Wiley-Liss, Inc.     

Thalamic input to inferior area 6 and area 4 in the macaque monkey

Recent cytoarchitectonic, histochemical, and hodological studies in primates have shown that area 6 is formed by three main sectors: the supplementary motor area, superior area 6, which lies medial to the spur of the arcuate sulcus, and inferior area 6, which is located lateral to it. Inferior area 6 has been further subdivided into two histochemical areas: area F5, located along the inferior limb of the arcuate sulcus, and area F4, located between area F5 and area 4 (area F1). The present study traced the thalamocortical projections of inferior area 6 and the adjacent part of area 4 by injecting small amounts of WGA-HRP in specific sectors of the agranular frontal cortex. Our data showed that each histochemical area receives a large projection from one nucleus of the ventrolateral thalamus (motor thalamus) and additional projections from other nuclei of this thalamic sector. Area F5 receives a large projection from area X of Olszewski ('52) and additional projections from the caudal part of the nucleus ventralis posterior lateralis, pars oralis (VPLo), and the nucleus ventralis lateralis, pars caudalis (VLc) (VPLo-VLc complex). Area F4 receives a large projection from the nucleus ventralis lateralis, pars oralis (VLo), and additional projections from area X and the VPLo-VLc complex. The rostral part of area F1 is innervated chiefly by VLo, plus smaller contributions from rostral VPLo and the VPLo-VLc complex. The caudal part of F1 receives its greatest input from VPLo, with a small contribution from VLo. In addition, each histochemical area receives projections originating from the intralaminar thalamic nuclei, the posterior thalamus, and--for area F4 and area F5--also from the nucleus medialis dorsalis (MD). Analysis of the physiological properties of the various histochemical areas in relation to their main thalamic input showed that those cortical fields in which distal movements are predominant (area F5, caudal part of area F1) are innervated chiefly by area X and VPLo, whereas those cortical fields in which proximal movements are predominant receive their main input from VLo. Because VPLo and area X are targets of cerebellothalamic pathways, whereas VLo receives a pallidal input, we propose that the cortical fields in which distal movements are most heavily represented are mainly under the influence of the cerebellum, whereas the cortical fields in which proximal movements are most heavily represented are mainly under the influence of the basal ganglia.     

Connections of the retrosplenial dysgranular cortex in the rat.

Although the retrosplenial dysgranular cortex (Rdg) is situated both physically and connectionally between the hippocampal formation and the neocortex, few studies have focused on the connections of Rdg. The present study employs retrograde and anterograde anatomical tracing methods to delineate the connections of Rdg. Each projection to Rdg terminates in distinct layers of the cortex. The thalamic projections to Rdg originate in the anterior (primarily the anteromedial), lateral (primarily the laterodorsal), and reuniens nuclei. Those from the anteromedial nucleus terminate predominantely in layers I and IV-VI, whereas the axons arising from the laterodorsal nucleus have a dense terminal plexus in layers I and III-IV. The cortical projections to Rdg originate primarily in the infraradiata, retrosplenial, postsubicular, and areas 17 and 18b cortices. The projections arising from visual areas 18b and 17 predominantly terminate in layer I of Rdg, axons from contralateral Rdg form a dense terminal plexus in layers I-IV, with a smaller number of terminals in layers V and VI, afferents from postsubiculum terminate in layers I and III-V, and the projection from infraradiata cortex terminates in layers I and V-VI. The efferent projections from Rdg are widespread. The major cortical projections from Rdg are to infraradiata, retrosplenial granular, area 18b, and postsubicular cortices. Subcortical projections from Rdg terminate primarily in the ipsilateral caudate and lateral thalamic nuclei and bilaterally in the anterior thalamic nuclei. The efferent projections from Rdg are topographically organized. Rostral Rdg projects to the dorsal infraradiata cortex and the rostral postsubiculum, while caudal Rdg axons terminate predominantely in the ventral infraradiata and the caudal postsubicular cortices. Caudal but not rostral Rdg projects to areas 17 and 18b of the cortex. The Rdg projections to the lateral and anterior nuclei also are organized along the rostral-caudal axis. Together, these data suggest that Rdg integrates thalamic, hippocampal, and neocortical information.     

Afferent and efferent innervation patterns of the superior olivary nucleus of the leopard frog

An HRP study of the frog's superior olivary nucleus (SON) revealed that it (1) receives reciprocal tonotopic projections from dorsal medullary nuclei and principal nuclei of the torus semicircularis (TS) bilaterally; (2) receives bilateral projections from caudalis nuclei and brainstem reticular nuclei, and unilateral projections from the ipsilateral ventral tegmental nuclei and the contralateral SON; (3) is reciprocally connected with the ipsilateral laminar and magnocellular nuclei of the TS, dorsal tegmental nuclei and the posterior thalamic nucleus; (4) projects directly to the ipsilateral central thalamic nucleus.     

Multiple topographically organized projections connect the central nucleus of the inferior colliculus to the ventral division of the medial geniculate nucleus in the gerbil, Meriones unguiculatus

The ventral division of the medial geniculate nucleus (MGv) receives almost all of its ascending input from the ipsilateral central nucleus of the inferior colliculus (CNIC). In a previous study (Cant and Benson [2006] J. Comp. Neurol. 495:511-528), we made injections of biotinylated dextran amine into the CNIC of the gerbil and demonstrated that it can be divided into two parts. One part (zone 1) receives almost all of its ascending input from the cochlear nuclei, the nuclei of the lateral lemniscus, and the main nuclei of the superior olivary complex; the other part (zone 2) receives inputs from the cochlear nuclei and nuclei of the lateral lemniscus but few or no inputs from the main olivary nuclei. Here we show that these two parts of the CNIC project differentially to the MGv. Axons labeled anterogradely by injections in zone 1 project throughout the rostral two-thirds of the MGv, whereas axons from zone 2 project to the caudal third of the MGv. Throughout much of their extent, the terminal fields do not appear to overlap, although both parts of the CNIC project to medial and dorsal parts of the MGv, and there may be overlap in the most ventral part as well. The results indicate that two parallel pathways arising in the CNIC remain largely separate in the medial geniculate nucleus of the gerbil. It seems most likely that the neurons in the two terminal zones in the MGv perform different functions in audition.     

Connections of the retrosplenial granular b cortex in the rat

Although the retrosplenial granular b cortex (Rgb) is situated in a critical position between the hippocampal formation and the neocortex, surprisingly few studies have examined its connections carefully. The present experiments use both anterograde and retrograde tracing techniques to characterize the connections of Rgb. The main cortical projections from Rgb are to the caudal part of the anterior cingulate cortex, area 18b, retrosplenial granular a cortex (Rga), and postsubiculum, and less dense terminal fields are present in the prelimbic and caudal occipital cortices. The major subcortical projections are to the anterior thalamic nuclei and the rostral pontine nuclei, and very small terminal fields are present in the caudal dorsomedial part of the striatum, the reuniens and reticular nuclei of the thalamus, and the mammillary bodies. Contralaterally, Rgb primarily projects to itself, i.e., homotypically, and more sparsely projects to Rga and postsubiculum. In general, the axons from Rgb terminate ipsilaterally in cortical layers I and III-V and contralaterally in layer V, with a smaller number of terminals in layers I and VI. Thalamic projections from Rgb target the anteroventral and laterodorsal nuclei of the thalamus, with only a few axons terminating in the anterodorsal nucleus, the reticular nucleus, and the nucleus reuniens of the thalamus. Rgb is innervated by the anterior cingulate cortex, precentral agranular cortex, cortical area 18b, dorsal subiculum, and postsubiculum. Subcortical projections to Rgb originate mainly in the claustrum, the horizontal limb of the diagonal band of Broca, and the anterior thalamic nuclei. These data demonstrate that, in the rat, Rgb is a major nodal point for the integration and subsequent distribution of information to and from the hippocampal formation, the midline limbic and visual cortices, and the thalamus. Thus, similarly to the entorhinal cortex, Rgb in the rat is a prominent gateway for information exchange between the hippocampal formation and other limbic areas of the brain.     

Connections of the bullfrog striatum: Efferent projections

Autoradiographic and degeneration techniques were used to describe striatal efferents in the bullfrog (Rana catesbeiana). Horseradish peroxidase (HRP) was then placed in the major terminal fields to reveal the striatal cells responsible for these projections. Except for the small ventral eminence of the lateral pallium immediately adjacent to the dorsal striatum, no pallial region receives a striatal projection. Most striatal efferents descend in the lateral forebrain bundle (LFB), passing through the anterior entopedun-cular nucleus, where one large fascicle decussates in the anterior commissure and innervates the contralateral anterior entopeduncular nucleus and caudal ventral striatum. A smaller fascicle exits the LFB to terminate in the ipsilateral lateral amygdala. The remaining efferents continue caudal in the LFB through the posterior entopeduncular nucleus, with sparse projections to the ventral thalamus, the adjacent preoptic areas, and the posterior tuberculum leaving the bundle at various points. At pretectal levels, some efferents leave the LFB to run dorsally, through the caudal pole of the central thalamic nucleus and into the posterior division of the lateral nucleus and the lateral portion of the posterior thalamic nucleus. Efferents also continue caudal, through the superficial tegmental cell groups (nucleus profundus mesencephali and superficial isthmal reticular nucleus) before turning dorsomedially into the ventral anterodorsal, lateral anteroventral, and rostral pole of the posterodorsal tegmental fields. A small superficial projection continues to isthmal levels but cannot be traced beyond. Tegmental HRP injections retrogradely fill cells in the dorsal and ventral striatum as well as nucleus accumbens and the anterior entopeduncular nucleus. Pretectal HRP injections fill cells only in the caudal ventral striatum and anterior entopeduncular nucleus. Anterior entopeduncular nucleus HRP injections fill numerous cells in all striatal divisions, but some of this filling may be due to interrupted fibers of passage. Thus the anuran striatum, which receives its major input from thalamic nuclei relaying tectal and toral input, can in turn influence the midbrain roof via several disynaptic pathways: through the anterior entopeduncular nucleus, pretectum, and tegmentum, all of which project directly to the tectum and torus (Wilczynski and Northcutt, ′77; Wilczynski, ′81). Additional trisynaptic routes through the anterior entopeduncular nucleus and its pretectal and tegmental connections parallel the striatal routes.     

Connections of the posterior thalamic region with the auditory ectosylvian cortex in the dog

The purpose of the present study was to define auditory cortical areas in the dog on the basis of thalamocortical connectivity patterns. Connections between the posterior thalamic region and auditory ectosylvian cortex were studied using axonally transported tracers: fluorochromes and biotinylated dextran amine. Cyto- and chemoarchitecture provided grounds for the division of the posterior thalamic region into three complexes, medial geniculate body (MGB), posterior nuclei (Po), and lateromedial and suprageniculate nuclei (LM-Sg). Distinctive cytoarchitectonic features and the distribution of dominant thalamocortical connections (determined quantitatively) allowed us to define four ectosylvian areas: middle (EM), anterior (EA), posterior (EP), and composite (CE). We found that each area was a place of convergence for projections from five to eleven nuclei of the three thalamic complexes, with dominant projections derived from one or two nuclei. Dominant topographical projections from the ventral nucleus to area EM confirmed physiological reports that it may be considered a primary auditory area (AI). We found the anterior part of the EM to be distinct in having unique strong connections with the deep dorsal MGB nucleus. Area EA, which receives dominant projections from the lateral Po (Pol) and medial MGB nuclei, as well as area EP, which receives dominant connections from the dorsal caudal MGB nucleus, compose two parasensory areas. Area CE receives dominant projections from the extrageniculate nuclei, anterior region of the LM-Sg, and Pol, supplemented with an input from the somatosensory VP complex, and may be considered a polymodal association area.