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PubMed1994

Movement- and task-related activations of motor cortical areas: a positron emission tomographic study.

Using repeated measurements of regional cerebral blood flow with positron emission tomography, we investigated the regional cortical activations induced in 10 normal subjects, by two different finger motor tasks, i.e., a repeated flexion-extension of all fingers and a repeated flexion-extension of the middle finger. The all-finger movement only activated the primary sensorimotor cortex (SM) and the supplementary motor area (SMA) contralateral to the movement. However, the activation of the SMA was clearly task related during this motor task, because it was only observed when the movement was triggered by an auditory cue but not when it was self-paced. The middle finger movement was performed during self-paced conditions. It induced a much more complex pattern of activation than the all-finger movement, characterized by a high degree of SM and SMA activation contralateral to the side of the movement, as well as a slight ipsilateral activation of these areas. We suggest that this pattern of cortical activation may reflect the process of individuating finger movement or the early stages of motor learning of this unusual and technically difficult movement. Our data also confirm that the SM activation is closely linked to the intrinsic parameters of the movement; while the SMA may be activated by different aspects of the movement realization and preparation.

Adult↗
PubMed1995

Increased sensitivity to ipsilateral cutaneous stimuli following transcranial magnetic stimulation of the parietal lobe.

Transcranial magnetic stimulation of the sensorimotor cortex results in decreased sensitivity of threshold electrical stimuli to fingers of the contralateral hand. It has been suggested that one factor contributing to neglect contralateral to a unilateral parietal lesion is a release of the normal hemisphere from reciprocal interhemispheric inhibition by the damaged hemisphere. Consistent with this account, the current study demonstrated that transcranial magnetic stimulation over the parietal cortex results in increased sensitivity to cutaneous stimulation ipsilateral to the stimulation. The likely mechanism is a transcranial magnetic stimulation-induced transient dysfunction of the ipsilateral parietal cortex that then results in disinhibition of the contralateral parietal cortex.

Adolescent↗
PubMed1997

Stereotactic thalamotomy in tremor-dominant Parkinson's disease: an H2(15)O PET motor activation study.

Stereotactic thalamotomy is an effective treatment for severe drug-resistant tremor. The thalamus, however, facilitates motor activity, and thalamotomy would be predicted to inhibit movement-associated cortical activation. Two tremulous parkinsonian patients were studied with H2(15)O positron emission tomography before and after left ventralis intermedius thalamotomy. Subjects were scanned at rest and during performance of externally paced joystick movements in freely selected directions with the right hand. Thalamotomy relieved tremor but, as predicted, led to decreased activation of the left sensorimotor cortex, lateral premotor cortex, and parietal area 7 on hand movement.

Aged↗
PubMed2001

Organization of corticostriatal projections from the vibrissal representations in the primary motor and somatosensory cortical areas of rodents.

To characterize corticostriatal projections from rodent sensorimotor cortex, the anterograde tracers biotinylated dextran amine (BDA) and fluororuby (FR) were injected into the whisker representations of the primary motor (MI) and somatosensory (SI) cortices. Reconstructions of labeled terminals and their beaded varicosities in the neostriatum and thalamus were analyzed quantitatively to determine the degree of labeled overlap in both of these subcortical structures. Corticostriatal projections from the vibrissal representation in MI were more extensive than corresponding projections from SI. Both cortical areas sent dense projections to the dorsolateral neostriatum, but the MI vibrissal representation also projected to regions located more rostrally and medially. Despite these differences, both MI and SI projected to overlapping parts of the dorsolateral neostriatum. Tracer injections in both cortical areas also produced dense anterograde and retrograde labeling in the medial sector of the posterior complex of the thalamus (POm). Because POm is somatotopically organized and has reciprocal connections with both SI and MI cortices, the amount of labeled overlap in POm was used to indicate whether the tracers were injected into corresponding whisker representations of MI and SI. We found that the proportion of labeled overlap in the neostriatum was highly correlated with the amount of labeled overlap in POm. These results indicate that the rodent neostriatum receives convergent projections from corresponding regions in MI and SI cortex. Furthermore, the thalamocortical projections of the POm indicate that it may modulate corticostriatal outputs from corresponding representations in MI and SI.

Animals↗
PubMed2004

The dorsolateral corticospinal tract in mice: an alternative route for corticospinal input to caudal segments following dorsal column lesions.

In rodents, the main contingent of corticospinal tract (CST) axons descends in the ventral part of the dorsal column. There is, however, a contingent of CST axons that descends in the dorsolateral column (the "dorsolateral corticospinal tract," or DLCST). Here, we define some of the features of the DLCST by tracing CST projections following injections of biotinylated dextran amine into the sensorimotor cortex, assessing the distribution of DLCST axons and terminal arborizations in intact mice and in mice in which the main contingent of CST axons in the dorsal column had been transected. Axons of the DLCST diverge from the main tract at the pyramidal decussation, gather in fascicles in the dorsolateral gray matter below the spinomedullary junction, and project in a gradual trajectory laterally toward the dorsolateral column over the first few cervical segments. DLCST axons then project along the dorsolateral column to sacral levels, giving rise to collaterals that project into the gray matter. Labeled DLCST axons were most abundant in cervical segments, where they were often collected in fascicles, and progressively decreased in number in more caudal segments. Tracing of DLCST axons in mice with selective lesions of the dorsal column revealed that DLCST axons arborize extensively throughout the dorsal and ventral horns and that the overall territory that the DLCST axons invade is similar to the territory innervated by the CST axons in the main tract. Some DLCST axon arbors with varicosities are seen near large neurons in the ventral horn (presumed motoneurons). Substantial numbers of DLCST axons project across the midline to the gray matter on the contralateral side. Thus, the DLCST provides an alternate route for CST input to caudal segments, which is of particular relevance for studies of CST distribution and function following partial spinal cord injuries.

Animals↗
PubMed2005

RhoA, RhoB, RhoC, Rac1, Cdc42, and Tc10 mRNA levels in spinal cord, sensory ganglia, and corticospinal tract neurons and long-lasting specific changes following spinal cord injury.

Inhibition of RhoA has been shown to enhance axonal regeneration following spinal cord injury. Here we mapped mRNA expression patterns of RhoA, B, and C, Rac1, Cdc42, and Tc10 in spinal cord, sensory ganglia, and sensorimotor cortex in uninjured rats, and following spinal cord injury or sham laminectomy. In the intact spinal cord, neurons displayed high levels of Rac1, Cdc42, and Tc10 mRNA hybridization signal. GFAP-immunoreactive astrocytes expressed primarily RhoB and Rac1, while oligodendrocyte-like cells expressed RhoA, Rac1, and Cdc42. Injury caused profound, long-lasting upregulation of RhoA, Rac1, Cdc42, and Tc10 mRNA in the spinal cord, while RhoB was modestly increased and RhoC did not change. GFAP-immunoreactive reactive astrocytes exhibited a dramatic increase of RhoA mRNA expression along with increases of Rac1 and Cdc42. Injury also led to elevation of RhoA, Cdc42, and Tc10 in neurons and modest increases of RhoA, Rac1, and Tc10 in oligodendrocyte-like cells. Laminectomy caused similar, but less pronounced alterations of investigated mRNA species. In dorsal root ganglia neuronal RhoA, Rac1, Cdc42, and Tc10 mRNA levels were increased similarly by spinal cord injury and sham surgery. The CST pyramidal cells expressed Tc10 mRNA and the CST itself was Tc10-immunoreactive. Tc10-immunoreactivity disappeared distal to injury. We conclude that there are gene-specific patterns of expression of the six different Rho-GTPases in normal spinal cord and dorsal root ganglia, and that specific changes of temporal and spatial expression patterns occur in response to spinal cord injury, suggesting different roles of these GTPases in the cellular sequelae of CNS injury.

Animals↗
PubMed1976

Cortical cells projecting to the dorsal column nuclei of cats. An anatomical study with the horseradish peroxidase technique.

The retrograde transport of horseradish peroxidase (HRP) has been used to label cortical neurons which give origin to descending projections to the dorsal column nuclei (DCN) in kittens and adult cats. HRP was injected unilaterally or bilaterally into the dorsal medulla at, or rostral to, the level of the obex, and, in most cases, retrogradely transported HRP was visualized in cortical neurons by incubation of serial 40 mum coronal or sagittal frozen sections. In the remaining cases, 1 mm coronal or sagittal slabs through the anterior part of the cerebral hemisphere were cut and incubated en bloc, and embedded in celloidin for sectioning (40 mum). HRP-positive neurons identifiable as the source of cortical projections to the DCN were layer V pyramidal cells whose largest diameter ranged from 20 to 45 mum: no "giant" cells were labelled. The cells appeared concentration in the fore-and hindlimb regions of the sensorimotor cortex and, to a lesser extent, in the second somatosensory area contralateral to the injected side. Labelled neurons were especially numerous in the upper bank of the cruciate sulcus and in the medial wall of the posterior sigmoid gyrus which respectively form parts of areas 4 and 3a (Hassler and Muhs-Clement, '64). The number of labelled neurons progressively diminished in the first somatosensory area proper (areas 3b, 1 and 2). Ipsilaterally, fewer labelled neurons were present, mainly in areas 4 and 3a. These results are in genral agreement with previous anatomcial and electrophysiological studies. It is suggested that previous results based upon lesions or electrical stimulation of the cerebral cortex in cats have failed to reveal adequately the cortical regions containing the bulk of the cells projectng to the DCN, since these regions have escaped selective experimental manipulation due to their relative inaccessibility.

Animals↗
PubMed1976

An anatomical study of some somesthetic afferents to the intercollicular terminal zone of the midbrain of the opossum.

Following lesions of the dorsal column nuclei, spinal cord and sensorimotor cortex, anterograde degeneration was traced to a common target in the central midbrain; the intercollicular terminal zone. Although not all the midbrain projections of the three pathways are contained within this zone, it receives overlapping projections from each of the three body-related somesthetic pathways studied. The intercollicular terminal zone covers the entirety of the external nucleus of the inferior colliculus and the intercollicular nucleus of Mehler, spilling over into adjacent parts of the central gray and deep layers of the superior colliculus. Little evidence of somatotopy in the somesthetic projections was disclosed, and bilateral input was found in each experimental case, regardless of the type of lesion. Golgi-stained material was examined to explore the basis of the extensiveness of the terminal zone. These materials showed that the dendritic spread of neurons in the external nucleus of the inferior colliculus and the intercollicular nucleus, taken together, covers the entirety of the intercollicular terminal zone. That is, the dendrites invade the same portions of the central gray and deep layers of the superior colliculus which are covered by the terminal field. We conclude that a tripartite division of the sensory midbrain may be justified: with the somesthetic modality being represented anatomically in a fashion at least roughly analogous to that of vision and audition.

Afferent Pathways↗
PubMed1981

Cortical and spinal somatosensory input to the superior colliculus in the golden hamster: an anatomical and electrophysiological study.

The horseradish peroxidase technique was used to identify the sources of somatosensory afferent fibers to the hamster superior colliculus. These experiments demonstrated that the tectum receives axons from pyramidal cells in layer V of the ipsilateral sensorimotor cortex, contralateral lamina IV of all levels of the spinal cord, the contralateral dorsal column nuclei, lateral cervical nucleus, internal basilar nucleus, and nucleus of the spinal trigeminal tract. Electrical stimulation of the spinal cord coupled with extracellular single unit recordings concentrated, for the most part, in the posterior portion of the tectum revealed that such stimuli activated approximately 40% of the cells tested. Almost all of these units were isolated ventral to the stratum opticum and 86% were responsive only to somatosensory stimulation. Analysis of the latencies of collicular responses obtained with two point spinal stimulation in intact hamsters and in animals subjected to somatosensory cortical and/or spinal damage indicated that the initial impulse elicited from most collicular cells was mediated by a polysynaptic pathway(s) which probably synapses in the dorsal column, lateral cervical, and/or internal basilar nuclei. Damage to the dorsal spinal cord and/or somatosensory cortex altered neither the incidence nor the response characteristics of spinally driven collicular neurons. This indicated that most somatosensory collicular cells also received input from the spinotectal fibers which travel in the ventrolateral quadrant. Electrical stimulation of somatosensory cortex activated about 20% of the cells tested in the ipsilateral superior colliculus. If cortical and spinal stimulation were delivered with an interstimulus interval ranging between 50 and 80 msec the response of the tectal neuron to the latter stimulus was suppressed in most cases. This was true regardless of the order of the stimulus pairing. Concurrent somatosensory cortical shocks also suppressed responses to tactile stimuli for 21% of the cells tested.

Afferent Pathways↗
PubMed1982

A light and electron microscopic study of regrowing pyramidal tract fibers.

Autoradiographic and EM techniques were used to study the regenerative capacity of severed axons in the mammalian CNS. In infant and adult hamsters the pyramidal tract was severed unilaterally in the medulla several millimeters rostral to the decussation. After survival to adulthood, the animals received injections of [3H] proline in the sensorimotor cortex ipsilateral to the lesion. Autoradiography showed that labeled pyramidal tract axons in the medulla did not cross the lesion site. Instead, in animals with infant lesions there was massive new axonal growth arising from the severed pyramidal tract several millimeters rostral to the cut. Most of these labeled fibers crossed to the contralateral brainstem, coalesced into a compact bundle, descended just medial to the spinal trigeminal nucleus, and grew caudally for 6-7 mm. Although the trajectory of the regrowing axons was completely abnormal, their pattern of termination in the dorsal column nuclei and dorsal horn of the cervical spinal cord was normal. Synapse formation by the anomalous regrowing pyramidal tract axons in their appropriate terminal areas was confirmed by electron microscopy of terminal degeneration in animals with infant pyramidotomies followed by adult cortical lesions. Autoradiographic labeling of the new pathway at short postlesion survival times showed that the fibers grew out rapidly at about 1 mm/day, a rate somewhat slower than normal (2-4 mm/day). There was a dramatic difference in the capacity of the pyramidal axons to regrow in animals operated as infants vs. those operated as adults. The regrowth was maximal with lesions at 4-8 days of age. Capacity for new growth declined sharply thereafter such that after 20 days of age, pyramidal tract lesions elicited no new growth but instead a progressive axon degeneration retrograde to the lesion. These results, in contrast to many previous findings, show that significant regrowth of severed axons can occur in the neonatal CNS. Most importantly pyramidal tract fibers regrowing by anomalous routes can nevertheless establish synaptic connections in appropriate terminal areas and thus, as we show in the following paper, play a functional role in maintaining normal motor behavior.

Age Factors↗
PubMed1983

Plasticity of the corticospinal tract following midthoracic spinal injury in the postnatal rat.

Rats received a midthoracic spinal cord "overhemisection" including right hemicord and left dorsal funiculus at birth (neonatal operates, N = 15) or 21 days of age (weanling operates, N = 14). In a second experiment neonatal (N = 6), 6-day (N = 3), and 12-day (N = 7) rats sustained a right sensorimotor cortex (SmI) ablation to destroy the left corticospinal tract (CST) at the same time as the spinal injury (double lesion operates). Later (3-12 months) injections of 3H-proline and autoradiography were used to label the left or right CST. The results of the first experiment showed that most right CST axons failed to grow around the spinal lesion in neonatal operates (N = 9). There was an increase in the density of label, mainly to CST projection areas, in a 1-mm zone rostral to the lesion. However, left CST axons bypassed the lesion by growing through the intact tissue in neonatal operates (N = 6). These displaced axons were consistently located within the dorsal portion of the lateral funiculus (dLF) and remained within that location caudal to the lesion, an area normally containing only a few CST axons. In spite of this abnormal position, these axons terminated bilaterally throughout the remainder of the cord in normal CST sites. In weanling operates, CST axons severed by the lesion did not regenerate around the lesion site. An increased density of label over the few spared axons within the left dLF and in CST projection zones immediately caudal to the lesion site suggested axonal sprouting by these axons. The results of the second experiment showed that the lack of growth of right CST axons around this injury in neonatal operates was, at least partially, due to an interaction with left CST axons. In neonatal double lesion operates, right CST axons grew around the spinal injury for a varying distance within the left dLF and distributed bilaterally to normal CST sites. The number of right CST axons bypassing the lesion was related to the configuration of the lesion site. A smaller number of right CST axons bypassed the lesion in 6-day double lesion operates and most terminated within 2-3 mm of the lesion site. Right CST axons failed to grow around this injury in 12-day double lesion operates.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1987

Ultrastructural analyses of afferent terminals in the subthalamic nucleus of the cat with a combined degeneration and horseradish peroxidase tracing method.

The synaptic organization of the feline subthalamic nucleus (STN) was studied electron microscopically. Following horseradish peroxidase (HRP) injections into the globus pallidus (GP) and electrolytic lesions of the nucleus tegmenti pedunculopontinus pars compacta (TCP) in the same cat, both degenerating and HRP-labeled terminals were found in the STN with abundant retrogradely HRP-labeled neurons. Degenerating terminals of TPC origin were medium-sized and characterized by asymmetric synaptic contacts. They synapsed widely on the STN neuronal surface, including the somata, dendrites of varying dimensions, dendritic spines and vesicle-containing processes. They formed 25.1%, 65.1%, 4.7%, and 4.7%, respectively, of all TPC efferent terminals. Some of the postsynaptic components were labeled with HRP. Occasionally both degenerating terminals and HRP-labeled terminals were in synaptic contact with the same HRP-labeled neuron: therefore, afferents of TPC and GP converge on the same STN projection neuron. In order to discover the origin of these HRP-labeled terminals, a mixed solution containing HRP and kainic acid was injected into the GP. Numerous degenerating terminals were observed to synapse with HRP-labeled STN neurons, but no HRP-labeled terminal was observed. These degenerating terminals were similar in appearance to the above-mentioned HRP-labeled terminals. They were characterized by their relatively large size, predominantly symmetric synapses, and preferential distribution on the somata and large or medium-sized dendrites. They formed 39.6%, 20.1%, and 31.1%, respectively, of all GP efferent terminals. Therefore, it became clear that both the HRP-labeled terminals of the first experiment and the degenerating terminals of the second experiment originated from the GP. Following surgical ablations of the primary sensorimotor cortex (Cx), some axon terminals in the STN showed degeneration. These degenerating terminals were small and formed asymmetric synapses mainly with dendritic spines, small dendrites and vesicle-containing processes. They formed 48.0%, 28.0%, and 12.0%, respectively, of all Cx efferent terminals. These electron microscopic investigations reveal the convergence of TPC and GP afferents and that STN projection neurons relay the TPC and pallidal inputs directly to the GP.

Afferent Pathways↗
PubMed1988

GABA-immunoreactive synaptic boutons in the rat basal forebrain: comparison of neurons that project to the neocortex with pallidosubthalamic neurons.

Although the basal forebrain, including the globus pallidus, contains a high concentration of gamma-aminobutyric acid (GABA), it is not known whether all types of neuron in the globus pallidus receive GABAergic synaptic input. We have studied two types of neuron: typical pallidal neurons that project to the subthalamic nucleus and magnocellular neurons which are found in the medial and ventral borders of the globus and project to the sensorimotor cortex. The postembedding immunogold staining of endogenous GABA revealed many preterminal axons and synaptic boutons that contained GABA immunoreactivity. Neurons that projected to the neocortex were postsynaptic to some of the GABA-immunoreactive boutons, the majority of which formed symmetrical membrane specializations. From a series of random electron micrographs through the perikarya and proximal dendrites of such retrogradely labelled neurons the density of GABA-containing afferent synaptic boutons was estimated to be 0.58 GABA-containing boutons per 100 micron of neuronal membrane. The GABA-containing boutons accounted for 72% of the total afferent input in the proximal regions of the pallidocortical neurons examined. The pallidosubthalamic neurons received many more afferent boutons than did the cortically projecting neurons, a high proportion (80.4%) of which were immunoreactive for GABA. The density of GABA-containing boutons in contact with pallidosubthalamic neurons was 8.9 boutons per 100 micron. It is concluded that cortically projecting basal forebrain neurons, that are probably cholinergic, are innervated by GABA-containing afferent boutons. However, pallidosubthalamic neurons in the same part of the basal forebrain are much more densely innervated by GABA-containing boutons.

Animals↗
PubMed1989

Glutamate and aspartate immunoreactivity in corticospinal neurons of rats.

A combination of retrograde tracers and immunostaining was employed to test whether corticospinal tract (CST) neurons in rats may use amino acid excitatory neurotransmitters. CST neurons were identified following injections of either Diamidino Yellow (DY) or colloidal gold-labeled enzymatically inactive horseradish peroxidase conjugated to wheat germ agglutinin (WGAapoHRP-Au) in the spinal cord. As retrograde tracers, the two substances seemed to be equally effective, but WGAapoHRP-Au was better suited than DY as a tracer to use in combination with immunocytochemistry. Sections through the primary sensorimotor cortex, which contained the bulk of identified CST neurons, and the secondary somatosensory cortex were processed with antisera raised in rabbits against glutamate (Glu) or aspartate (Asp) conjugated by glutaraldehyde to hemocyanin. In rats with DY injections, about 60-75% of the CST neurons were Glu-immunopositive, with higher ratios in SI and MI than in SII. Similar results were obtained in all areas examined from the rats with injections of WGAapoHRP-Au. Only sections from rats with injections of WGAapoHRP-Au were processed for Asp immunostaining. In this material, between 65 and 75% of the CST neurons were Asp-immunopositive, with a slightly higher ratio in SI and MI than in SII. The possibility that these results might reflect limited penetration of the antiserum and/or staining of the same population of CST neurons by either antiserum was addressed in sections processed with both the Glu and Asp antisera. In sections incubated in a mixture of the two antisera, the percentage of immunostained CST neurons was higher, about 90%, than in sections processed for only one of the two antisera. Furthermore, in rats in which Glu and Asp antibodies were visualized by two distinguishable immunostainings, four populations of CST neurons were identifiable: 1) neurons only immunopositive for Glu, 2) neurons only immunopositive for Asp, 3) neurons likely to be stained by both, and 4) neurons immunonegative for both antisera. Twenty-five to 30% of CST neurons were positive for only one antiserum, and about 50% were positive for both. No preferential distribution was evident for any one of these populations of neurons. However, perikaryal cross-sectional areas were larger for the double-stained than for the single-stained CST neurons. Glutamergic and aspartergic transmission in CST neurons has been proposed in several publications in which methods other than immunocytochemistry were employed.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1990

Branching patterns of corticospinal axon arbors in the rodent.

Despite extensive study of corticospinal connections in a variety of species, little is known about the detailed morphology of corticospinal axon arbors. Results in previous studies of primates based on intra-axonal filling with horseradish peroxidase (HRP) staining of a limited sample of fibers suggest that corticospinal arbors branch widely to multiple motoneuronal pools. To determine whether this pattern of corticospinal connectivity is present in nonprimate species as well, we studied the branching patterns of corticospinal axon arbors in a rodent species, the golden hamster. The axons were labeled by iontophoretic injection of Phaseolus vulgaris-leucoagglutinin (PHA-L) into small regions of the forelimb and hindlimb sensorimotor cortex, and immunohistochemistry with the peroxidase-antiperoxidase (PAP) method was used to reveal fine details of terminal arbors within the cervical and lumbar enlargements of the spinal cord. As in higher mammals, corticospinal connections are topographically organized. Moreover, corticospinal axons arising from somatosensory cortex project primarily to the dorsal horn, whereas those from motor cortex terminate most heavily in the ventral horn. This differential projection pattern, not previously demonstrated in rodents, implies functional differences between somatosensory and motor components of the corticospinal pathway. Reconstruction of corticospinal arbors in the ventral horn showed that in both cervical and lumbar spinal cord segments, axons branch widely into interneuronal regions. A surprising number appear to extend into motoneuron cell groups, and some of these axons branch into multiple motoneuronal pools. Widely divergent corticospinal axons that branch to multiple motoneuron pools have been shown to mediate activity in functionally related muscle groups of the primate forearm. The present results suggest that in other species, such as the rodent, a similar divergence of corticospinal arbors may also function to facilitate activity in subsets of muscles.

Animals↗
PubMed1990

Specificity of corticospinal axon arbors sprouting into denervated contralateral spinal cord.

Previous studies have reported considerable plasticity in the rodent corticospinal pathway in response to injury. This includes sprouting of intact axons from the normal pathway into the contralateral spinal cord denervated by an early corticospinal lesion. We carried out the present study to obtain detailed information about the time course, origin, and degree of specificity of corticospinal axons sprouting in response to denervation. Hamsters (Mesocricetus auratus) ranging in age from 5 to 23 days received unilateral lesions of the left medullary pyramidal tract. Two weeks after the lesion, small regions of the right sensorimotor cortex opposite the lesion were injected with the plant lectin Phaseolus vulgaris leucoagglutinin (PHA-L). After a further 2 week survival period, immunohistochemistry was carried out on frozen sections of the fixed brains and spinal cords. Detailed morphological analysis of PHA-L labeled corticospinal axons revealed that sprouting from the intact corticospinal pathway into the contralateral denervated spinal cord occurred only at local spinal levels and not at the pyramidal decussation. Arbors sprouting into the denervated cord frequently arose from corticospinal axons that branched into the normal side of the cord as well. Sprouting was maximal after early lesions (5 days) and declined with lesions at later ages up to 19 days. Sprouting corticospinal axons arborized with the same degree of functional and topographic specificity as previously reported for normal corticospinal arbors (Kuang and Kalil: J. Comp. Neurol. 292:585-598, '90), such that axons arising from somatosensory cortex projected only to the dorsal horn, those from motor cortex innervated only the ventral horn, and normal forelimb and hindlimb topography was preserved. Sprouting fibers also had normal branching patterns. Parallel studies of developing corticospinal arbors showed that sprouting could not be attributed to maintenance or expansion of early bilateral connections. These results suggest that local signals, most likely similar to those governing normal corticospinal development, elicit corticospinal sprouting and determine specificity of axon arbors.

Aging↗
PubMed1991

Corticomotoneuronal connections in the rat: evidence from double-labeling of motoneurons and corticospinal axon arborizations.

In order to investigate the possibility of direct corticomotoneuronal (CM) connections in the rat, an anterograde-retrograde double-labeling method was developed. Phaseolus vulgaris-leucoagglutinin (PHA-L) anterograde tracing of corticospinal axons was combined with retrograde labeling of spinal motoneurons either by a conjugate of choleragen subunit B with horseradish peroxidase (CB-HRP) or by wheat germ agglutinin (WGA). The location of PHA-L injection unilaterally in the forelimb area of sensorimotor cortex and the CB-HRP or WGA injections in corresponding contralateral wrist or digit extensors or flexors were determined and matched on the basis of movement responses elicited by intracortical microstimulation. Light microscopic observation showed, in addition to the main contralateral dorsal corticospinal tract (CST), the presence of four other CST minor components in the contralateral lateral, ipsilateral ventral, and ipsilateral dorsal funiculi of the cervical spinal white matter and at the base of contralateral dorsal horn of the gray matter, respectively. PHA-L-labeled CST axonal arbors were observed from Rexed's lamina I through lamina X of contralateral spinal gray matter, most extensively in laminae VI and VII; some CST axons reached the zone of motoneuronal somata in lamina IX and a few of them also entered the lateral and occasionally the ventral funiculi, ramifying in the white matter. Between the zones of PHA-L-labeled CST axonal arbors on the one hand and CB-HRP/WGA labeled spinal motoneuronal somata with their extensive dendritic trees on the other, there was a large overlap, covering partly both the gray and the white matter. PHA-L-labeled axonal boutons (en passant or terminaux) were seen to contact the dendrites or even the somata of motoneurons in the gray matter, according to light-microscopic criteria for identification of synaptic contacts. Axodendritic CM contacts were occasionally observed in the lateral funiculus of the white matter as well. In general, only a single contact was observed between an individual PHA-L-labeled CST axon and a given retrogradely labeled motoneuron. In contrast to the common notion that direct CM connections are a specialty of primates, the present morphological data support the presence of direct CM connections also in some other mammals, such as the rat.

Afferent Pathways↗
PubMed1994

Immunohistochemical detection of calcium/calmodulin-dependent protein kinase II in the spinal cord of the rat and monkey with special reference to the corticospinal tract.

Calcium/calmodulin-dependent protein kinase II is a prominent enzyme in the mammalian brain that phosphorylates a variety of substrate proteins. In the present study, monoclonal antibodies that specifically recognize either the alpha or the beta isoforms of this enzyme were used to determine the distribution of these isoforms within the rat and monkey spinal cord. In the rat, the corticospinal tract consists of two components: the dorsal corticospinal tract, which occupies the ventralmost aspect of the dorsal funiculus; and the ventral corticospinal tract, which occupies an area adjacent to the ventral median fissure. Both dorsal and ventral corticospinal tract fibers were strongly immunopositive for the alpha-antibody. Unilateral ablation of the sensorimotor cortex of the rat eliminated the alpha-immunoreactive staining in the contralateral dorsal corticospinal tract. The neuropil in the superficial laminae of the dorsal horn (Rexed's laminae I and II) was densely stained with the alpha-antibody, whereas the neuropil in laminae IV-X was immunonegative. Dense alpha-immunopositive neurons were also distributed in the head of the dorsal horn (laminae I-IV). In contrast to the strong alpha-immunoreactivity seen in the dorsal corticospinal tract fibers, only very weak beta-immunoreactivity was observed in this tract. Moderate beta-immunoreactive products were distributed homogenously throughout the neuropil of the gray matter, although the neuropil of the superficial laminae of the dorsal horn (laminae I and II) was stained more strongly than the other regions of the gray matter (laminae III-X). Neuronal components in all laminae were immunopositive for the beta-antibody. Thus, motoneurons in the ventral horn, which were immunonegative for the alpha-antibody, were immunopositive for the beta-antibody. This selective distribution pattern of immunoreactivity of alpha- and beta-antibodies in the rat was also present in the monkey spinal cord, although the alpha-immunopositive corticospinal tract fibers in the monkey descended in the lateral funiculus as the lateral corticospinal tract instead of passing through the dorsal funiculus, as is the case in the rat. The differential distribution of immunoreactivity in the spinal cord suggests that these two isoforms of calcium/calmodulin-dependent protein kinase II may have different functional roles in the spinal cord.

Animals↗