Role of the commissural channels of the somatosensory system in the conduction of feedback signals during the control of voluntary movements

The time-course of formation of the conditioned defence reflex and electromyograms of the working forelimb were studied on cats with classical and commissural somatosensory pathways transections. It was established that exclusion of the classical somatosensory projections only reduces the rate of the skill formation that is related to the possibility of conducting feedback signals via the remaining commissural pathways. However, the commissural channel in question is not effective enough, since its isolated functioning is accompanied by an increase in the duration of instrumental movement. The combined transection of the classical and commissural somatosensory pathways excludes the possibility of formation of the conditioned reflex on the "deafferented" limb. The lack of direct visual control over the moving forelimb does not essentially affect the behavioral and electromyographic data.     

Bilateral receptive fields in cortical area SII: contribution of the corpus callosum and other interhemispheric commissures

The corpus callosum contributes to the interhemispheric transfer of somatosensory information. Since the somatosensory pathways are essentially crossed, a number of studies have postulated that the corpus callosum may be responsible for the presence of bilateral receptive fields (RFs) in cortical area SII. Moreover, subcortical structures, as well as some of the other commissures, may also contribute to the bilateral nature of these cells. In order to assess the relative importance of the corpus callosum, this study compared the RF properties of cells in area SII of callosum-sectioned cats to normal cats, using single-cell recordings. Results showed that the corpus callosum makes an important contribution to the bilateral activation of cells in SII, since the proportion of cells with bilateral RFs found in callosum-sectioned cats was less than half that obtained in normal cats. The decrease in the proportion of bilateral RFs was found for all body regions with the exception of the face. However, the substantial number of bilateral RFs remaining in callosotomized cats indicates that this structure is not the sole contributor to the bilateral activation of cells in SII. In order to determine whether this residual bilateral activation might be mediated by the other interhemispheric commissures, a group of cats was subjected, besides the callosotomy, to the additional transection of their subcortical commissures, including the anterior, posterior, habenular, and intertectal commissures, as well as the massa intermedia. When this group of deep-split cats was compared to the callosotomized group, the results indicated that the contribution of the other commissures to bilateral activation is negligible, since approximately the same proportion of bilateral RFs was encountered in the two groups. The relative importance of the callosal contribution to bilateral RFs of different body regions is discussed with respect to the roles commonly attributed to this structure.     

A possible principle of kinesthetic compensation

The movement of the cat forelimb was studied under the conditions of complete transection of the classical somatosensory pathways from the limb to the cortex of the contralateral hemisphere. It was demonstrated that in animals with the intact system of the somatosensory (kinesthetic) feedback, the parameters of the same memorized movement varied widely from response to response whereas the exclusion of lemniscus projections from the control system led to the poor movement pattern and to the appearance of motor stereotypy. The hypothesis is advanced that deficit of kinesthetic information in this case is compensated for not only by the other intact connections of the kinesthetic analyzer but also due to the conversion of the nervous system to the central-programmed level control which itself demands less participation of the somatosensory feedback. The non-crossed projections of the kinesthetic analyzer are capable of forming and correcting this program and of ensuring the effect afferentation as well. The same principle of the functioning may take place in any compensatory reconstructions in the CNS.     

Role of the neocortical and subcortical commissures in the interhemispheric transfer of visual information to the hippocampus

The light flash evoked potentials were recorded in the hippocamp of cats with transected left optic tract and cerebral commissures. The data obtained show a significant reduction in the response amplitude (by 1.5-2 times) in both hemispheres after unilateral transection of the optic tract. The minimal response amplitude (10-15 times less than normal) was recorded in the left hippocamp of cats with additional transection of the commissures of the endbrain, mesencephalon and diencephalon. In contrast, the latent period of the first and second components of the evoked potential was similar to that in the control group animals. It is suggested that the neocortical commissural visual inputs play the most important role in interhemispheric transmission of visual information to the hippocamp.     

Commissural channels of visual information transmission

Experiments were made in adult cats with different transections of the classical and commissural visual tracts to study evoked potentials and unit neuronal activity in response to visual stimulation. The commissural channels of the telencephalon, diencephalon and mesencephalon were demonstrated to be highly effective in visual information conduction to the visual cortex. Complete transection of the classical and commissural tracts with the exception of the commissural tract of the third ventricle fundus and midbrain reticular formation resulted in the disappearance of evoked potentials in the visual cortex in response to light flashes, despite the fact that 8 of the 15 recorded neurons responded to photostimulation. Activation of such neurons was caused by visual information conduction via two possible commissural channels: midbrain reticular formation and subventricular interhemispheric commissures of the diencephalon.     

Maze behavior in macaques

Specialization of the right cerebral hemisphere in man for analysis of spatial relations suggests that a similar specialization may exist in macaques. This possibility is being examined in animals first trained to negotiate a maze and then subjected to transection of optic chiasm and forebrain commissures. In the one successful experiment to date, clear evidence of such specialization was obtained for the left hemisphere in a monkey consistently using its left hand for retrieval of food. Provisional evidence for the existence of unilateral engrams in such circumstances was also obtained by limiting input during training to one eye and hemisphere and testing the other before and after completion of transection of the forebrain commissures. Finally, it was found that in the split brain macaque, a lesion in the parietal-occipital area of one hemisphere initially disrupted maze performance using either hemisphere.     

Changes in Choline Acetyltransferase Activity and High-Affinity Choline Uptake,but Not in Acetylcholinesterase Activity and Muscarinic Cholinergic Receptors,in Rat Somatosensory Cortex after Sciatic Nerve Injury

Selected cholinergic markers (choline acetyltransferase, acetylcholinesterase, muscarinic acetylcholine receptor, high-affinity choline uptake) were studied in the hindlimb representation areas of the rat somatosensory cortex and within the visual cortex 1 to 63 days after unilateral transection of the sciatic nerve. In the contralateral somatosensory cortex, peripheral deafferentation resulted in a significant reduction of choline acetyltransferase activity (by 15%) 3 days after sciatic nerve injury, and in a significant reduction of high-affinity choline uptake (by 30%) 1 day after nerve transection, in comparison to untreated control rats. Investigations in individual cortical layers revealed that the decrease of both choline acetyltransferase activity and high-affinity choline uptake sites was mainly due to reductions in cortical layer V. Acetylcholinesterase activity and [³H]quinuclidinyl benzilate binding to muscarinic acetylcholine receptors were not affected by unilateral transection of the sciatic nerve. In the ipsilateral somatosensory cortex, as well as in the visual cortex at both cortical hemispheres, no significant changes in the cholinergic parameters studied could be detected. The data indicate that peripheral deafferentation of the somatosensory cortex results in a transient change of presynaptic cholinergic parameters within the affected somatosensory area as early as 1 to 3 days after the lesion; thus, they emphasize the involvement of cholinergic mechanisms in cortical reorganizational events.     

Changes in choline acetyltransferase activity and high-affinity choline uptake, but not in acetylcholinesterase activity and muscarinic cholinergic receptors, in rat somatosensory cortex after sciatic nerve injury

Selected cholinergic markers (choline acetyltransferase, acetylcholinesterase, muscarinic acetylcholine receptor, high-affinity choline uptake) were studied in the hindlimb representation areas of the rat somatosensory cortex and within the visual cortex 1 to 63 days after unilateral transection of the sciatic nerve. In the contralateral somatosensory cortex, peripheral deafferentation resulted in a significant reduction of choline acetyltransferase activity (by 15%) 3 days after sciatic nerve injury, and in a significant reduction of high-affinity choline uptake (by 30%) 1 day after nerve transection, in comparison to untreated control rats. Investigations in individual cortical layers revealed that the decrease of both choline acetyltransferase activity and high-affinity choline uptake sites was mainly due to reductions in cortical layer V. Acetylcholinesterase activity and [3H]quinuclidinyl benzilate binding to muscarinic acetylcholine receptors were not affected by unilateral transection of the sciatic nerve. In the ipsilateral somatosensory cortex, as well as in the visual cortex at both cortical hemispheres, no significant changes in the cholinergic parameters studied could be detected. The data indicate that peripheral deafferentation of the somatosensory cortex results in a transient change of presynaptic cholinergic parameters within the affected somatosensory area as early as 1 to 3 days after the lesion; thus, they emphasize the involvement of cholinergic mechanisms in cortical reorganizational events.     

Chronological observations of primary somatosensory cortical maps in kittens following low thoracic (T12) spinal cord transection at 2 weeks of age.

The present study investigated the reorganization of the somatosensory cortex in kittens following T12 spinal cord transection at 2 weeks of age. Multiunit electrophysiological methods were used to map the somatosensory cortex of kittens at 3, 6, and 9 weeks after the transection. The entire reorganized cortical region was driven by substitute cutaneous inputs, primarily from the trunk, at 3 weeks after spinal cord transection. Although the level of cortical responsiveness remained the same throughout the 9 weeks studied, internal trunk representation changed, and there was an increase in shoulder girdle representation and emergence of forelimb representation. Poor somatotopic and topographic order was observed in the reorganized cortex, regardless of time posttransection. Finally, trunk receptive fields displayed a wide variety of shapes, sizes, and orientations not seen in the normal cortex.     

Chronological Observations of Primary Somatosensory Cortical Maps in Kittens following Low Thoracic (T12) Spinal Cord Transection at 2 Weeks of Age

The present study investigated the reorganization of the somatosensory cortex in kittens following T₁₂ spinal cord transection at 2 weeks of age. Multiunit electrophysiological methods were used to map the somatosensory cortex of kittens at 3, 6, and 9 weeks after the transection. The entire reorganized cortical region was driven by substitute cutaneous inputs, primarily from the trunk, at 3 weeks after spinal cord transection. Although the level of cortical responsiveness remained the same throughout the 9 weeks studied, internal trunk representation changed, and there was an increase in shoulder girdle representation and emergence of forelimb representation. Poor somatotopic and topographic order was observed in the reorganized cortex, regardless of time posttransection. Finally, trunk receptive fields displayed a wide variety of shapes, sizes, and orientations not seen in the normal cortex.     

Muscarinic cholinergic receptor binding in rat hindlimb somatosensory cortex following partial deafferentation by sciatic nerve transection.

Peripheral nerve injury or amputation leads to extensive changes within the central representations of the mammalian body surface. The mechanisms responsible for post-traumatic reorganization of these maps in adults may also, at least partly, underlie a more general feature of the somatosensory system--the capacity for stimulus-dependent plasticity. Acetylcholine has been implicated in both of these processes. We studied the binding of the ligands [3H]QNB and [3H]pirenzepine in rat hindlimb somatosensory cortex from 1 to 14 days following sciatic nerve transection. Although the [3H]QNB binding was not different from normal levels in tissue homogenates of the affected somatosensory cortex, differences were demonstrated when binding was measured on a layer-by-layer basis. [3H]QNB binding was changed only in certain layers, at certain times. The predominant effects appeared to be a decrease in binding in the middle layers from 4 to 14 days after the transection. Combining the [3H]QNB data with data obtained from the more M1-selective ligand [3H]pirenzepine suggested that complex changes occur among several muscarinic receptors, including receptors with non-M1 subtype characteristics. Moreover, unilateral nerve transection affects the hindlimb somatosensory regions in both hemispheres.     

Neuroanatomy of the terminal (sixth abdominal) ganglion of the crayfish,Procambarus clarkii (Girard)

Summary We studied the neuroanatomy of the terminal (sixth abdominal) ganglion in the crayfish Procambarus clarkii with silver-impregnated sections and nickel fills. We describe the fiber tracts, commissures and neuropilar areas, and give the topological relationships of motoneurons and intersegmental interneurons with reference to their neuropilar landmark structures.All five anterior abdominal ganglia have an almost identical number of 600–700 neurons with a similar pattern of distribution. Each contains a single neuromere with a common plan of neuropil organization. In contrast, the terminal ganglion consists of two neuromeres which appear to be derived from the intrinsic sixth abdominal and telson ganglion. The basic organization of each neuromere parallels that of the third abdominal ganglion in the appearance and arrangement of fiber tracts and commissures, although some modifications occur. The fusion of two neuromeres is represented by the duplication of segmentally homologous neurons, MoGs and LGs, whose topological relationships to the neuropil structures are similar to those of the anterior ganglion.We also discuss the origin of the telson and its ganglion (the seventh abdominal neuromere), and dispute the classical theory that the telson derives from a postsegmental region.     

Behavioral plasticity and central regeneration of locomotor reflexes in the freshwater oligochaete, Lumbriculus variegatus. I: Transection studies

Abstract. Access to the ventral nerve cord in living specimens of Lumbriculus variegatus , an aquatic oligochaete, is normally impossible because surgical invasion induces segmental autotomy (self-fragmentation). We show here that nicotine is a powerful paralytic agent that reversibly immobilizes worms, blocks segmental autotomy, and allows experimental access to the nerve cord. Using nicotine-treated worms, we transected the ventral nerve cord and used non-invasive electrophysiological recordings and behavioral analyses to characterize the functional recovery of giant nerve fibers and other reflex pathways. Initially, after transection, medial giant fiber (MGF) and lateral giant fiber (LGF) spikes conducted up to, but not across, the transection site. Reestablishment of MGF and LGF through-conduction across the transection site occurred as early as 10 h (usually by 20 h) after transection. Analyses of non-giant-mediated behavioral responses (i.e., helical swimming and body reversal) were also made following nerve cord transection. Immediately after transection, functional reorganization of touch-evoked locomotor reflexes occurred, so that the two portions of the worm anterior and posterior to the transection site were independently capable of helical swimming and body reversal responses. Similar reorganization of responses occurred in amputated body fragments. Reversion back to the original whole-body pattern of swimming and reversal occurred as early as 8 h after transection. Thus, functional restoration of the non-giant central pathways appeared slightly faster than giant fiber pathways. The results demonstrate the remarkable plasticity of locomotor reflex behaviors immediately after nerve cord transection or segment amputation. They also demonstrate the exceptional speed and specificity of regeneration of the central pathways that mediate locomotor reflexes.     

Coordination between locomotor and respiratory rhythms in the great ramshorn snail Planorbarius corneus: Transmitter-dependent modifications

In a reduced preparation of Planorbarius corneus consisting of the CNS and mantle complex, both the dopamine precursor L-DOPA and the serotonin precursor 5-HTP have been found to be able to induce and maintain rhythmic pneumostome (PN) movements phase-coupled to fictive cyclic locomotion in a neurotransmitter-specific manner. After the transection of pedal commissures, pharmacologically induced PN movements were coordinated with the locomotor activity rhythm generated by the left pedal ganglion, as in Lymnaea regardless of spatial inversion of its CNS. Nevertheless, in Planorbarius during the 5-HTP-induced fictive muscular locomotion, the PN was never opened, but cuddled up to the mantle at the same phase of the locomotor cycle corresponding to close down the PN in Lymnaea.     

The nervous system in the praesoma of Echinorhynchus salmonis (Acanthocephala: Echinorhynchidae)

Based on light microscopic observations, nerve pathways are described for the first time in the praesoma of a species of the Echinorhynchidae. The pathways are described for 18 nerves, 8 paired and 2 single, which originate from the cerebral ganglion and a post-ganglionic cell and terminate in the body wall musculature and the proboscis. The location of two commissures formed by these nerves is also described.     

Age-related dynamics of the parameters of somatosensory evoked potentials in healthy children

The purpose of this study was to evaluate the parameters of somatosensory evoked potentials (SSEPs) in healthy children of different age groups (n = 94). The amplitudes of the main cortical peaks and the central sensory conduction time (CSCT) from n. medianus and n. tibialis in children aged under 12 months, 1–12 years, and 12–17 years were estimated and compared. No significant cortical peaks were recorded from the tibial nerve in five children younger than 1 year (5 out of 23, 22%). Significant differences in CSCT were observed between the children younger than 1 year and two other groups. The amplitudes did not significantly differ between the groups. Thus, SSEPs may be used for the evaluation of somatosensory pathways in children aged one month to 17 years. CSCT differs significantly between children younger than 1 year and other age groups. Age-related reduction in CSCT and elevation of the cortical peak amplitudes may reflect the myelination of somatosensory pathways and the improvement in nervous system integration.     

Nonoverlapping Thalamocortical Connections to Normal and Deprived Primary Somatosensory Cortex for Similar Forelimb Receptive Fields in Chronic Spinal Cats

The fluorescent dye retrograde tracing technique, using fast blue in combination with fluorogold, was used to examine thalamocortical projections from the ventrobasal complex to primary somatosensory cortex in chronic spinal cats that sustained T12 cord transection at 2 weeks of age. Following cord transection at this age, it has been shown that forelimb afferents can excite the deprived hindlimb projection zone, in addition to the region of somatosensory cortex that they normally occupy (McKinley et al, 1987). These two regions of cortex are separated by over 10 mm, thus facilitating the determination of whether the forelimb representation in “hindlimb cortex” is derived from the sector of the ventrobasal complex of the thalamus representing the forelimb, hindlimb, or both. Injections of the two dyes into separate regions of the cortex that were excited by the same peripheral forelimb receptive fields produced single labeling of two nonoverlapping clusters of thalamic neurons. This finding suggests that the projections for these two areas are independent and distinct, and indicates that altered thalamocortical projections do not contribute the critical component underlying reorganizational changes observed at the cortical level after spinal cord transection. It is hypothesized that the degree of reorganization required to achieve the magnitude of change observed in the cortex must occur below the level of the thalamocortical relay.     

Sensory projections of identified coxal hair sensilla of the scorpionHeterometrus fulvipes (Scorpionidae)

The topography of long hair sensilla on the coxae of walking legs and pedipalps of the scorpionHeterometrus fulvipes is described. Identified long hair sensilla are cobalt filled, and central projections of sensory fibres are reported for the first time in the suboesophageal ganglion of this scorpion. The afferent fibres arising from each long hair sensilla segregate into ventral, dorsomedial and dorsal tracts upon their entry into the suboesophageal ganglion. These transverse tracts bifurcate towards the middle of the leg neuromere and form three ipsilateral, plurisegrnental, longitudinal sensory pathways. Filling a pair of bilaterally distributed long hair sensilla shows bilaterally arranged longitudinal afferent tracts interconnected by distinct transverse commissures. Similar patterns of sensory projections are observed when filling homologous hairs on other legs and pedipalps. Numerous fine collaterals arise from the longitudinal sensory trancts that subdivide and end in small blebs presumed to be presynaptic endings. The dorsal and dorsomedial longitudinal tracts and their respective commissures are in close association with the dendritic arborisations of pedipalpal and leg motor neurons, suggesting direct contact between them. The probable functions of these multisegmental hair afferent pathways are discussed.     

Transneuronal induction of the highly sialylated isoform of the neural cell adhesion molecule following nerve injury

The polysialylated, embryonic form of the neuronal cell adhesion molecule (PSA-NCAM) is known to participate in a whole series of synaptic rearrangements even in adult animals. The possible role of this molecule in neuroplastic changes of the adult rat somatosensory cortex induced by unilateral transection of the infraorbital branch of the trigeminal nerve was studied with PSA-NCAM immunostaining at light microscopic level. Two- and three-month-old CFY albino rats were sacrificied on days 1, 4, 6, 14 and 21 following operation and PSA-NCAM immunoreaction was examined at three levels of the vibrissa-cortex neuraxis, namely, in the principal nucleus of the trigeminal nerve, in the ventral posteromedial nucleus of the thalamus and in the somatosensory cortex. The lower levels of the neuraxis remained free of PSA-NCAM labeling, similarly to control, intact animals. However, a large number of scattered small neurons became PSA-NCAM immunoreactive in layers IV-VI on both ipsi- and contralateral sides of the somatosensory cortex from day 6 onwards, suggesting a possible transynaptic regulation of NCAM sialylation state.     

Anatomy of neurons crossing the tritocerebral commissures of the cockroach Periplaneta americana (Blattaria)

The neuronal connections of the tritocerebral commissures of Periplaneta americana were studied in the brain-suboesophageal ganglion complex and the stomatogastric nervous system by means of heavy metal iontophoresis through cut nerve ends followed by silver intensification. The tritocerebral commissure 1 (Tc1) contains mainly the processes of the subpharyngeal nerve (Spn) whose neurons are located in both tritocerebral lobes and in the frontal ganglion. Some neurons of the frontal ganglion project through the Tc1 to the contralateral tritocerebrum. A few fibers in this commissure were observed projecting to the protocerebrum and the suboesophageal ganglion. There are tritocerebral neurons which pass through the Tc1 or the tritocerebral commissure 2 (Tc2) and extend on into the stomatogastric nervous system. One axon of a descending gaint neuron appears in the Tc2. This neuron lies in the tritocerebrum and connects the brain to the contralateral side of the ventral nerve cord. In addition, sensory fibers of the labral nerve (Ln) traverse both commissures to the opposite tritocerebrum. The anatomical and physiological relevance of the identified neuronal pathways is discussed. © 1995 Wiley-Liss, Inc.