Spatial relationships between the terminations of somatic sensory and motor pathways in the rostral brainstem of cats and monkeys. I. Ascending somatic sensory inputs to lateral diencephalon

Projections to the lateral diencephalon from the dorsal column nuclei (DCN), lateral cervical n. (LCN), and spinal cord (ST) in cats and monkeys, and from the spinal portion of the trigeminal n. (sTN) in the cat were compared using a double-orthograde labeling strategy. This strategy combines autoradiographic and degeneration tracing methods in the same animal and permits direct comparisons of the terminal labeling patterns of two different pathways in each experiment. The results suggest that the major part of the lateral diencephalon which receives input from the somatic sensory pathways in both the cat and the monkey is arranged in a core-shell fashion. The core consists of the group of nuclei which together constitute the ventrobasal complex (VB). The shell consists of a group of nuclei which together tend to surround VB nearly completely. This group includes the posterior group (PO), the ventral posteroinferior n. (VPI), and the border region between VB and the ventrolateral n. (VB-VL). In addition to the core and shell regions, two other regions in the lateral diencephalon receive input from the somatic sensory pathways. These regions are the ventromedial part of the magnocellular portion of the medial geniculate n. (MGNm) and caudomedial portion of the zona incerta (ZI). The cytoarchitectural and hodological patterns of the core region differ from those of the shell region. In both the monkey and the cat, the core region (VB) has a relatively homogeneous cytoarchitectural appearance and is filled by dense inputs from DCN, LCN, and sTN in the cat and from DCN, LCN, and ST (and probably from sTN) in the monkey. Direct comparisons of the terminals of fibers from different pathways demonstrate that although there is some convergence on the same neurons within VB, the major tendency is for each of the inputs to form its densest terminations on different neurons. This partial segregation manifests itself in two ways. First, each pathway has its own preferred territory within VB where its terminations are the densest. Second, the terminal fields of the inputs usually have a clustered appearance which is characterized by dense patches of terminals separated by regions in which the terminations appear quite sparse. The dense patches from different pathways do not occur in relation to the same groups of neurons. In contrast, most portions of the shell region have a lower cell density than that of the core and a heterogeneous cytoarchitectonic appearance which can often be described as transitional in character between its neighboring areas. In both species, different parts of the shell region receive sparse and scattered input from those pathways which project densely and precisely to areas immediately adjacent to that part of the shell. Very few of the terminals of these different inputs appear to converge on the same groups of neurons. The two otehr recipient targets of somatic senory input (i.e., MGNm, ZI) each has its own characteristic connective pattern that differs from taht of either the core or the shell region. The connective patterns in the cat and monkey are quite similar. The mian differences are in the projections of parts of the ST ans LCN pathways. The nature of these differences suggest taht it might be useful from a functional perspective to consider the LCN and ST pathways together as part of the same spinal system, rather than as separate functional entities. The LCN pathyway could then be viewed as having perhaps been dervied from different parts of a single population of diencephalic-projecting neurons in the spinal cord of the two species. When these anatomical results are considered together with the available electrophysiological evidence, it appears that the response properties and functions of some portions of the somatic senory regions within the diencephalon can be generally predicted from knowledge of the particular pathways whose axons terminate within these regions. Such predictions can be made, however, only when the input pathways have markedly different functions (e.g., vestibular, auditory, cutaneous). At present, more precise kinds of predicitions are precluded by the similarity that exists between the functional properties of many of the units in teh DCN, sTN, LCN, and ST pathways, and the luck of knowledge of the sorting processes which occur as fibers in each of these pathways diverge to terminate in different parts of the brain.     

Organization of subcortical pathways for sensory projections to the limbic cortex. II. Afferent projections to the thalamic lateral dorsal nucleus in the rat

Afferent projections to the thalamic lateral dorsal nucleus were examined in the rat by the use of retrograde axonal transport techniques. Small iontophoretic injections of horseradish peroxidase were placed at various locations within the lateral dorsal nucleus, and the location and morphology of cells of origin of afferent projections were identified by retrograde labeling. For all cases examined, subcortical retrogradely labeled neurons were most prominent in the pretectal complex, the intermediate layers of the superior colliculus, and the ventral lateral geniculate nucleus. Labeled cells were also seen in the thalamic reticular nucleus and the zona incerta. Within the cerebral cortex, labeled cells were prominent in the retrosplenial areas (areas 29b, 29c, and 29d) and the presubiculum. Labeled cells were also seen in areas 17 and 18 of occipital cortex. Peroxidase injections in the dorsal lateral part of the lateral dorsal nucleus result in labeled neurons in all of the ipsilateral pretectal nuclei, but especially those that receive direct retinal afferents. Labeled cells were also seen in the ventral lateral geniculate nucleus and the rostral tip of laminae IV-VI of the superior colliculus. In contrast, peroxidase injections in ventral medial portions of the lateral dorsal nucleus result in fewer labeled pretectal cells, and these labeled cells are found exclusively in the pretectal nuclei that do not receive retinal afferents. Other labeled cells following injections in the rostral and medial portions of the lateral dorsal nucleus are seen contralaterally in the medial pretectal region and nucleus of the posterior commissure, and bilaterally in the rostral tips of laminae IV and V of the superior colliculus. Camera lucida drawings of HRP labeled cells reveal that projecting cells in each pretectal nucleus have a characteristic soma size and dendritic branching pattern. These results are discussed with regard to the type of sensory information that may reach the lateral dorsal nucleus and then be relayed on to the medial limbic cortex.     

Lamina I spinocervical tract terminations in the medial part of the lateral cervical nucleus in the cat.

The terminations of spinocervical tract fibers in the lateral cervical nucleus (LCN) of the cat were examined with anterogradely transported Phaseolus vulgaris leucoagglutinin (PHA-L) in order to analyze their organization relative to the most medial part and the main body (the lateral two-thirds) of the LCN, which have differential projections and physiological characteristics. Iontophoretic injections of PHA-L in laminae I-V of the spinal dorsal horn yielded dense labeling in somatotopically appropriate regions of the main body of the LCN, and, as seen previously with horseradish peroxidase, additional terminations were present in the medial LCN after injections at either cervical or lumbar spinal levels. The morphological characteristics of the PHA-L labeling in these two parts of the LCN were different. Terminations in the lateral LCN consisted of dense clusters of thick fibers bearing large numbers of boutons. The terminal axons in the medial part of the LCN displayed a reticulated network of longitudinally oriented, fine fibers with well-spaced varicosities. Some of the fine fibers in the medial LCN appeared to be collaterals of thicker fibers that terminated in the lateral LCN. Injections of PHA-L that were restricted to lamina I resulted in terminal labeling only in the medial LCN. The labeling was more sparse than that observed in the medial LCN after larger dorsal horn injections but displayed the same morphological characteristics. Lamina I terminations were seen in the medial LCN after cervical or lumbar injections on both the ipsilateral and contralateral sides. The PHA-L observations were corroborated by the presence of many retrogradely labeled lamina I cells at both cervical and lumbar spinal levels, following injections of cholera toxin subunit b or rhodamine-labeled microspheres in the medial LCN. In addition, double-immunofluorescent labeling for PHA-L and substance P was performed in a few cases, since substance P immunoreactivity is present in fibers in the medial LCN and also in cell bodies in lamina I; however, very few spinocervical fibers displayed immunoreactivity for both antigens. These observations indicate that the medial part of the LCN receives input from lamina I neurons, and probably from lamina III-V neurons as well, at cervical and lumbar spinal levels. The lamina I input to the medial LCN provides a basis for the small population of nociceptive neurons that differentiate the medial LCN. The lamina I input could also be responsible for the general inhibition of lateral LCN neurons by wide-field noxious stimulation, via activation of GABAergic interneurons in the medial LCN.(ABSTRACT TRUNCATED AT 400 WORDS)     

Organization of subcortical pathways for sensory projections to the limbic cortex. I. Subcortical projections to the medial limbic cortex in the rat

Subcortical afferent projections to the medial limbic cortex were examined in the rat by the use of retrograde axonal transport of horseradish peroxidase. Small iontophoretic injections of horseradish peroxidase were placed at various locations within the dorsal and ventral cingulate areas, the dorsal agranular and ventral granular divisions of the retrosplenial cortex and the presubiculum. Somata of afferent neurons in the thalamus and basal forebrain were identified by retrograde labeling. Each of the anterior thalamic nuclei was found to project to several limbic cortical areas, although not with equal density. The anterior dorsal nucleus projects primarily to the presubiculum and ventral retrosplenial cortex; the anterior ventral nucleus projects to the retrosplenial cortex and the presubiculum with apparently similar densities; and the anterior medial nucleus projects primarily to the cingulate areas. The projections from the lateral dorsal nucleus to these limbic cortical areas are organized in a loose topographic fashion. The projection to the presubiculum originates in the most dorsal portion of the lateral dorsal nucleus. The projection to the ventral retrosplenial cortex originates in rostral and medial portions of the nucleus, whereas afferents to the dorsal retrosplenial cortex originate in caudal portions of the lateral dorsal nucleus. The projection to the cingulate originates in the ventral portion of the lateral dorsal nucleus. Other projections from the thalamus originate in the intralaminar and midline nuclei, including the central lateral, central dorsal, central medial, paracentral, reuniens, and paraventricular nuclei, and the ventral medial and ventral anterior nuclei. In addition, projections to the medial limbic cortex from the basal forebrain originate in cells of the nucleus of the diagonal band. Projections to the presubiculum also originate in the medial septum. These results are discussed in regard to convergence of sensory and nonsensory information projecting to the limbic cortex and the types of visual and other sensory information that may be relayed to the limbic cortex by these projections.     

Functional organization of the ventral lateral geniculate complex of the tree shrew (Tupaia belangeri): II. Connections with the cortex,thalamus, and brainstem

Connections of the ventral lateral geniculate complex (GLv) in the tree shrew were traced by anterograde and retrograde transport of WGA-HRP. The results buttress earlier findings that GLv in this species is composed of two main divisions, lateral and medial, each of which differs in its connections with the brainstem and cerebral cortex. The connections of the lateral division (GLv) suggest that it participates in visuosensory functions: it receives input from the retina, striate cortex, pretectum, and retino-recipient layers of the superior colliculus. These connections help clarify the identification of the internal and external subdivisions of GLv inasmuch as projections from both the superior colliculus and pretectum terminate in the external subdivision and each, in turn, receives a projection from the internal subdivision. Connections of the medial division suggest that this part of the nucleus is involved with visuomotor functions. Thus, the medio-caudal subdivision projects to the pontine nuclei, the prerubral field and the central lateral nucleus. The medio-caudal subdivision also receives projections from the lateral cerebellar nucleus, so that the GLv-ponto-cerebello-GLv loop involves mainly one subdivision of GLv. The medio-rostral subdivision receives projections from the pretectum and parietal cortex. Its output is directed primarily at the intermediate and deep layers of the superior colliculus. All of these targets of GLv, the pons, prerubral field, and deep layers of the superior colliculus, are known to play a role in the coordination of head and eye movements. Additional connections of GLv with the vestibular nuclei, intralaminar nuclei, hypothalamus, and facial motor nucleus are also described. © 1993 Wiley-Liss, Inc.     

Avian somatosensory system: II. Ascending projections of the dorsal column and external cuneate nuclei in the pigeon

The ascending projections of the dorsal column and external cuneate nuclei (DCN/CuE) in the pigeon were investigated in anterograde tracing experiments by using autoradiography or wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP). The results show that the majority of ascending projections decussate via internal arcuate fibers to form a contralateral medial lemniscus which ascends in a ventral position. In the brainstem, terminal fields were observed in the ventral lamella of the inferior olive (OI), the parabrachial nuclei (PB) of the dorsolateral pons, the intercollicular nucleus (ICo) of the midbrain, and the nucleus pretectalis diffusus (PD). In the diencephalon there were terminal fields in the strata cellulare externum and internum (SCE and SCI) of the caudal hypothalamus; in the intercalated (ICT), ventrolateral (VLT), and reticular nuclei of the ventral thalamus; in the nuclei principalis precommissuralis (PPC), spiriform medialis (SpM), and dorsolateralis posterior, pars caudalis (cDLP) of the caudal thalamus; and in the nuclei dorsalis intermedius ventralis anterior (DIVA), dorsolateralis posterior, pars rostralis (rDLP), dorsolateralis anterior (DLA), and dorsolateralis anterior, pars medialis (DLM) of the rostrodorsal thalamus. The origins of these projections within the DCN/CuE complex were verified in retrograde tracing experiments with WGA-HRP and were found to be partly differentiable with respect to their targets. The projections to DIVA, rDLP, DLA, DLM, cDLP, and SpM arise from all rostrocaudal levels of the DCN/CuE complex; those to ICo arise from caudomedial nuclear regions, while those to the hypothalamus and ventral thalamus arise from rostrolateral nuclear regions. Projections to PB arise from lamina I neurons of the dorsal horn of upper cervical spinal cord segments and from CuE. No evidence was found of a projection to the cerebellum. The distribution of the cells of origin of the medial lemniscus (ML) within the DCN/CuE complex was found to be largely coextensive with the areas of termination of primary spinal (Wild: J. Comp. Neurol. 240:377-395, '85) and some trigeminal (Dubbledam and Karten: J. Comp. Neurol. 180:661-678, '78) afferents. Furthermore, the areas of termination of the ML within the rostrodorsal and caudal thalamus are also either coextensive or closely associated with nuclei which provide a somatosensory projection to separate regions of the telencephalon (Wild: Brain Res. 412:205-223, '87). There are thus clear similarities in the overall pattern of somatosensory projections in the pigeon and in many mammalian species.     

Modification of the projection from the sensory cortex to the motor cortex following the elimination of thalamic projections to the motor cortex in cats

Examination of the projection from area 2 of the sensory cortex to the motor cortex revealed substantial changes following lesion of the ventrolateral nucleus of the thalamus. These observed changes were as follows. (1) The polarity of the evoked potentials elicited by area 2 stimulation reversed in the depth of the motor cortex whereas in normal animals, there was no reversal. (2) The amplitude of area 2-elicited EPSPs in the motor cortical neurons became greater following the lesion of VL. (3) The shape of the observed EPSPs was characterized by multiple peaks whereas in normal animals, the EPSPs were generally smooth and monophasic. (4) Neurons receiving a short-latency input from area 2 were distributed throughout the depths of the motor cortex whereas in normal animals, they were located only in the upper layers (layers II and III). (5) Intracellular injection of HRP revealed that the neurons receiving short-latency input were not restricted to typical stellate type cells, but also included bipolar or bitufted neurons with elongated cell bodies and polarized arborizations. These neurons were located in the superficial (II and III) as well as in the deep (V) layer. It is concluded that the elimination of thalamic input resulted in the reinforcement of the corticocortical input to the motor cortex. The subsequently observed corticocortical projection extended to neurons did not originally innervated by the association fibers. The results suggested that functional recovery following thalamic lesion is partly due to reorganization of projections from the sensory cortex to the motor cortex.     

Substance P innervation of the rat and cat thalamus. II. Cells of origin in the spinal cord.

Evidence in the preceding paper suggests that fibers and terminals immunopositive for substance P (SP) in somatosensory thalamic nuclei are part of the spinothalamic tract (STT). In this paper, more direct evidence on this point is provided by immunocytochemistry for SP on the cervical spinal cord, alone or combined with the retrograde transport of colloidal gold-labeled wheat germ agglutinin conjugated to enzymatically inactive horseradish peroxidase (WGAapoHRP-Au). In cats and rats pretreated with colchicine and/or anterolateral chordotomy (to increase SP content in cell bodies), many small to large cell bodies are SP-immunopositive especially in laminae I and V, but also in more ventral laminae of the upper cervical cord. SP neurons are also present in the dorsolateral funiculus (in the lateral spinal nucleus, LSN, in rats) but not in the lateral cervical nucleus or in the internal basilar nucleus. In both species there is a considerable degree of overlap in the distribution of SP-positive neurons and that of STT neurons. SP immunocytochemistry in rats after WGAapoHRP-Au injection in the somatosensory thalamus reveals SP-positive STT neurons in LSN, in lamina I and in lamina V, and, to a lesser extent, in more ventral laminae. These results demonstrate that SP is a marker and/or neuromediator for some STT neurons. Together with the evidence discussed in the preceding paper, the results also suggest that SP-positive neurons may be involved in the transmission of nociceptive input.     

Topography and collateralization of the dopaminergic projections to motor and lateral prefrontal cortex in owl monkeys.

The sources and histochemical characteristics of dopaminergic projections to motor and premotor areas of cortex were investigated in owl monkeys in which information from related studies was used to subdivide cortex into motor fields. Brainstem projections to frontal cortex were identified by injections of different fluorescent dyes in the primary motor cortex (M1) and the supplementary motor area (SMA), first identified by microstimulation. Injections were also placed in dorsal premotor cortex and lateral prefrontal cortex. The distribution of retrogradely labeled neurons was related to the location of tyrosine hydroxylase immunolabeled neurons on the same or alternate brain sections to identify the dopamine (DA) neurons. All DA cortically projecting neurons were located in the A8-A10 complex, largely in its dorsal components, including the parabrachial pigmented n. of the ventral tegmental area (VTA), pars gamma of the substantia nigra compacta, and the dorsal part of the retrorubral area (A8). Fewer cells were in the midline groups of VTA (n. linearis rostralis and caudalis) and in the n. paranigralis. DA neurons projecting to M1, SMA, and prefrontal cortex were largely intermixed, and some of these neurons were double or triple labeled by the fluorescent dyes, indicating collateralization to two or three fields; DA cells projecting to M1 were more numerous than to the other locations. The dorsal components of the A8-A10 complex from which arose the DA mesocortical projection were also characterized by the presence of calbindin-immunoreactive neurons and by a dense neurotensin and noradrenergic terminal innervation. Compared to rodents or felines, the DA neurons projecting to the lateral frontal lobe of primates appear to be shifted dorsally and laterally in the nigral complex. The topographic overlap, partial collateralization, and common histochemical characteristics of the DA mesocortical neurons projecting to different fields of the lateral frontal lobe suggest that some degree of functional unity exists within this projection.     

Anatomical and physiological properties of the projection from the sensory cortex to the motor cortex in normal cats: the difference between corticocortical and thalamocortical projections

Details of the distribution of terminal sites of the projection fibers from area 2 of the sensory cortex to the motor cortex were studied and compared with the distribution of terminals from the ventrolateral (VL) nucleus of the thalamus to the motor cortex. The results obtained were as follows: Intracortical microstimulation (ICMS) in area 2 produced measurable short-latency EPSPs only in neurons located in layers II and III of the motor cortex, whereas VL stimulation produced short-latency EPSPs in neurons throughout the depths of the motor cortex. The time from the beginning to the peak of the EPSPs was not significantly different for area 2- and VL-elicited EPSPs suggesting that there was no systematic difference between effective terminal sites for both inputs. However, there was a difference when a given neuron received both inputs suggesting that there was a segregation between the two inputs within a given cell. The majority of area 2-elicited EPSPs were smooth and monophasic, but some (40%) of them showed double peaks indicating that some neurons received mono- and disynaptic inputs from area 2. Intracellular injections of HRP suggested that neurons receiving input from area 2 were predominantly multipolar non-pyramidal neurons in layers II and III whereas neurons receiving thalamic input were pyramidal as well as non-pyramidal cells. Field potentials in the motor cortex evoked by area 2 stimulation did not change polarity in the depths of the cortex and therefore, differed from the VL-evoked potentials suggesting differences in the mechanisms of generating the electrical fields. It is concluded that association fibers effective for producing EPSPs terminate primarily on non-pyramidal cells in layer II and III whereas VL fibers terminate not only on pyramidal but also on non-pyramidal cells in layers III and V. This study provided a basis for examining the modifiability of association fibers after elimination of VL input to the motor cortex which is reported in the following paper.     

Distribution of zebrin II in the gigantocerebellum of the mormyrid fish Gnathonemus petersii compared with other teleosts.

Immunocytochemistry has demonstrated unexpected heterogeneity among cerebellar Purkinje cells. For example, monoclonal antibody Mab anti-zebrin II reveals parasagittal bands of immunoreactive Purkinje cells in the mammalian cerebellum, but reveals a non-sagittal cerebellar compartmentation pattern in goldfish and gymnotiform fish. The present paper investigates the cerebellar compartmentation pattern, as reflected in the zebrin II distribution, in two other teleosts, the electric mormyrid fish Gnathonemus petersii with its large and regularly built gigantocerebellum, and the electrosensory osteoglossomorph teleost Xenomystis nigri, by using light as well as electron microscopic immunohistochemical techniques. Zebrin II is expressed only in Purkinje cells, where it is present in the cytoplasm of all neuronal compartments, including spines, distal and proximal dendrites, the cell body, and the initial part, as well as terminal boutons of the axon. Other types of cerebellar neurons, including the eurydendroid projection neurons, are zebrin II-negative. In Gnathonemus, zebrin II-positive Purkinje cells are present in the large caudolateral part of the valvula, in lobes C2, C3, and C4 of the corpus, and in the anterior as well as the posterior part of the caudal cerebellar lobe. Zebrin II-negative Purkinje cells are present in a continuous region encompassing the rostromedial part of the valvula, the lobus transitorius, lobe C1 and the ventral part of lobe C2, and in a small, lateral zone of the posterior part of the caudal lobe. In Xenomystis, all Purkinje cells, including those in the medial valvula and the posterior part of the caudal lobe, appear to react with mab anti-zebrin II. This more widespread distribution may be due to the presence of a second antigenic polypeptide in this species. On the basis of the present findings, it is concluded that the mormyrid lobus transitorius, lobe C1, and the ventral part of lobe C2 probably belong to the valvula, while the corpus is restricted to the dorsal part of lobe C2, lobe C3, and lobe C4. The functional significance of zebrin II expression for different subsets of teleostean Purkinje cells remains unclear, since comparisons of different teleosts reveal no general correlation with particular afferent or efferent connections, nor with special morphological features such as a dendritic palisade pattern or different arrangements of the Purkinje cell bodies. A comparison between mammals and teleosts suggests that a distinct parasagittal cerebellar zonation in teleosts is absent, and the major part of the teleostean cerebellum may be considered as a single (midsagittal) cerebellar zone, with about the same width as one mammalian parasagittal zone.     

Retinal projections to the superior colliculus and dorsal lateral geniculate nucleus in the tammar wallaby (Macropus eugenii): II. Topography after rotation of an eye prior to retinal innervation of the brain

Retinal projections to visual centers in a marsupial mammal, the tammar wallaby (Macropus eugenii), have been investigated after an eye rotation prior to retinal innervation of the brain. Retinal topography to the superior colliculus and dorsal lateral geniculate nucleus was mapped by using laser lesions of the retina and horseradish peroxidase histochemistry. Despite the change in orientation of optic axon outgrowth from the developing eye after rotation, retinal ganglion cells made orderly connections in the colliculus and geniculate according to their original retinal position within the eye and not their rotated position. Axons must have corrected their pathways at some point between the back of the eye and their targets. The optic chiasm was one such site. Optic axons from the rotated eye took an abnormal course at the caudal end of the chiasm. Growth of optic axons through aberrant pathways in the brain did not preclude specific innervation of targets. When by chance optic axons entered through the oculomotor nerve root they specifically innervated their correct visual centers, albeit in reduced density, and did not innervate inappropriate targets. These results support the idea of specific interactions between growing axons, the pathways they grow along, and their targets.     

Origins of callosal projections to the supplementary motor area (SMA): a direct comparison between pre-SMA and SMA-proper in macaque monkeys.

The two subdivisions of the supplementary motor area (SMA), the pre-SMA (rostrally) and SMA-proper (caudally), exhibit distinct functional properties and clear differences with respect to their connectivity with the spinal cord, the thalamus, and other homolateral motor cortical areas. The goal of the present study was to establish in monkeys whether these subdivisions also differ with regard to their callosal connectivity. Two fluorescent retrograde tracers (Fast Blue and Diamidino Yellow) were injected in each animal, one in the pre-SMA and the second in the SMA-proper. Tracer injections in the pre-SMA or in SMA-proper resulted in significant numbers of labeled neurons in the opposite SMA, premotor cortex (PM), cingulate motor areas (CMA), and cingulate gyrus. Labeled neurons in M1 were rare, being observed only after injection in the SMA-proper. The two subdivisions of the SMA differed in the proportion of labeled neurons found across areas providing their callosal inputs. The SMA-proper receives about half of its callosal inputs from its counterpart in the other hemisphere (42-65% across monkeys). A comparable proportion of neurons was found in the pre-SMA after injection in the opposite pre-SMA (32-47%). The pre-SMA receives more callosal inputs from the rostral halves of the dorsal PM, the ventral PM, and the CMA than from their caudal halves. In addition, the pre-SMA, but not the SMA-proper, receives callosal inputs from the prefrontal cortex. The SMA-proper receives more callosal inputs from the caudal halves of the dorsal PM and ventral PM than from their rostral halves. The two subdivisions of the SMA receive callosal inputs from the same cortical areas (except the prefrontal cortex and M1), but they differ with respect to the quantitative contribution of each area of origin. In conclusion, quantitative data now support the notion that pre-SMA receives more transcallosal inputs than the SMA-proper.     

Fluorescent compounds as retrograde tracers compared with horseradish peroxidase (HRP). II. A parametric study in the peripheral motor system of the cat

Fluorescent compounds which are currently used as retrograde tracers were tested in the cat peripheral motor system and compared with horseradish peroxidase (HRP). The tracers were either injected into forelimb muscles or applied to the proximal end of transected forelimb nerves. The remaining muscles of the limb has been carefully denervated. Following intramuscular injection all fluorescent compounds labeled spinal cord motoneurons, the DAPI compounds labeled endothelial cells in addition. In the nerve application mode tracer positive motoneurons were only observed when propidium iodide (PI) and the DAPI compounds were used, whereas bisbenzimide (BB), nuclear yellow (NY) and primuline did not label any cells. The fluorescence of BB labeled motoneurons were predominantly located in the cytoplasma. NY positive motoneurons showed a different localization of the fluorescent label between the different neurons of the same motornucleus: in some neurons it was exclusively located in the nucleus, in others predominantly in the cytoplasma, in the majority in both compartments. The intracellular distribution of the BB and the NY label was independent of the pH of the fixation fluid. The fluorescent tracers labeled the motoneuronal cell columns in their complete rostrocaudal extent and in a position identical to the one obtained with HRP. However, some substances (PI, fast blue) labeled less motoneurons of a motornucleus than did HRP, none of the fluorescent tracers labeled more. The results are discussed under several aspects: use of the investigated fluorescent compounds as single tracers; use of several tracers in the same animal to map collateral projections of one and the same neuron; use of several tracers in the same animal to establish the topographical relation between several independent neuronal populations.     

Direct evidence for the link between monoaminergic descending pathways and motor activity:: II. A study with microdialysis probes implanted in the ventral horn of the spinal cord

In order to define precisely the relation between descending monoaminergic systems and the motor system, we measured in the ventral horn of spinal cord of adult rats the variations of extracellular concentrations of 5-HT, 5-HIAA, DA and MHPG. Measurements were performed during rest, endurance running on a treadmill, and a post-exercise period, with microdialysis probes implanted permanently for 45 days. We found a slight decrease in both 5-HT and 5-HIAA during locomotion with a more marked decrease during the post-exercise period compared to the mean of rest values. In contrast, the concentration of DA and MHPG increased slightly during the exercise and decreased thereafter. These results, when compared with those of a previous study, which measured monoamines in the spinal cord white matter [C. Gerin, D. Bécquet, A. Privat, Direct evidence for the link between monoaminergic descending pathways and motor activity: I. A study with microdialysis probes implanted in the ventral funiculus of the spinal cord, Brain Res. 704 (1995) 191–201], highlight the complex regulation of the release of monoamines that occurs in the ventral horn.     

Direct evidence for the link between monoaminergic descending pathways and motor activity. I. A study with microdialysis probes implanted in the ventral funiculus of the spinal cord

Monoaminergic projections to the spinal cord are involved in the modulation of motor, autonomic, and sensory functions. More specifically, the increase of electrical activity of serotonergic neurons in raphe obscurus has been correlated with locomotion in t treadmill-trained cats [Jacobs,B.L. and Fornal, C.,Trends Neurosci., 9 (1993) 346–352]. In order to test the direct correlation between locomotion and the release of monoamines, microdialysis probes were permanently implanted for 45 days into the ventral funiculus of the spinal cord (white matter) of adult rats. Eight days after implantation, these rats were subjected to an endurant exercise on a treadmill, and dialysis sessions were organized in such a way that microdialysate samples of 15 min duration were collected during pre-, per- and post-exercise periods. Measurements of serotonin, 5-hydroxyindoleacetic acid, dopamine and 3-methoxy-4-hydroxyphenylethylglycol concentration in the extracellular space showed significant increases during locomotion when compared with both pre- and post-exercise values. Histological analysis shows that serotonergic axons were present close to the dialysis probe. These results demonstrate that the implantation of a microdialysis probe in the ventral funiculus, close to a potential target of monoaminergic projections, is a suitable technique for the collection of neuromediators released during spontaneous running.     

Lesion-induced synapse reorganization in the hippocampus of cats: sprouting of entorhinal, commissural/associational, and mossy fiber projections after unilateral entorhinal cortex lesions, with comments on the normal organization of these pathways.

This study evaluates whether three forms of sprouting occur in the hippocampus of the cat following unilateral entorhinal cortex (EC) lesions: (1) sprouting of projections from the EC contralateral to the lesion; (2) sprouting of the commissural/associational system; and (3) sprouting of mossy fibers. Tract tracing techniques were used to define the normal organization of the entorhinal cortical projection system, the commissural/associational (C/A) systems, and the mossy fiber projections in normal cats. The same techniques were then used to evaluate whether there were changes in these projections in animals with long-standing unilateral EC lesions. The projections from the entorhinal cortex were evaluated autoradiographically following injections of 3H proline into the entorhinal area. The projections of the C/A system were traced using the Fink-Heimer technique after lesions of the hippocampal commissures, and by using autoradiographic techniques after injections of 3H proline into the hippocampus. The distribution of mossy fibers was evaluated using the Timm's stain. The results reveal that unilateral lesions of the EC in cats lead to the same sorts of sprouting that have been described in rats. There is: (1) an increase in the density of the crossed projection from the surviving EC to the contralateral dentate gyrus that had been deprived of its normal EC inputs; (2) an expansion of the terminal field of the C/A projection system into portions of the molecular layer of the dentate gyrus normally occupied by EC projections; and (3) an increase in supragranular mossy fibers in some animals. The mossy fiber sprouting was especially prominent when the lesions encroached upon the hippocampus. The studies also reveal additional details about the normal organization of hippocampal pathways in cats. The most important points are: (1) there is a crossed projection from the entorhinal cortex to the contralateral dentate gyrus; and (2) there is a complex laminar organization of the commissural and associational terminal fields in the molecular layer of the dentate gyrus that appears to be related to the point of origin of the projections along the septotemporal axis of the hippocampus. This heretofore unrecognized aspect of the laminar organization of C/A terminations has important implications for the temporal competition hypothesis, which has been advanced to account for the development of these afferent systems.