Abstract: We used functional magnetic resonance imaging to differentiate cerebral areas involved in two different dimensions of haptic shape perception: encoding and matching. For this purpose, healthy right-handed subjects were asked to compare pairs of complex 2D geometrical tactile shapes presented in a sequential two-alternative forced-choice task. Shape encoding involved a large sensorimotor network including the primary (SI) and secondary (SII) somatosensory cortex, the anterior part of the intraparietal sulcus (IPA) and of the supramarginal gyrus (SMG), regions previously associated with somatosensory shape perception. Activations were also observed in posterior parietal regions (aSPL), motor and premotor regions (primary motor cortex (MI), ventral premotor cortex, dorsal premotor cortex, supplementary motor area), as well as prefrontal areas (aPFC, VLPFC), parietal-occipital cortex (POC) and cerebellum. We propose that this distributed network reflects construction and maintenance of sensorimotor traces of exploration hand movements during complex shape encoding, and subsequent transformation of these traces into a more abstract shape representation using kinesthetic imagery. Moreover, haptic shape encoding was found to activate the left lateral occipital complex (LOC), thus corroborating the implication of this extrastriate visual area in multisensory shape representation, besides its contribution to visual imagery. Furthermore, left hemisphere predominance was shown during encoding, whereas right hemisphere predominance was associated with the matching process. Activations of SI, MI, PMd and aSPL, which were predominant in the left hemisphere during the encoding, were shifted to the right hemisphere during the matching. In addition, new activations emerged (right dorsolateral pre-frontal cortex, bilateral inferior parietal lobe, right SII) suggesting their specific involvement during 2D geometrical shape matching.

Abstract: Over the past 30 years, extensive research has been conducted in the field of cortical plasticity, under the impetus of seminal studies showing that the mature brain retains a capacity to reorganize the morphological and functional architecture of its neural circuits in order to adapt to environmental changes and mediate functional recovery following injury. Much effort has been focused on determining how idiosyncratic experience translates into molecular, structural and physiological changes in the sensory and motor representations embedded within cortical networks. The wealth of data generated by a broad spectrum of experimental manipulations has allowed unprecedented progress in our understanding of the physiological processes and neuroplasticity mechanisms underlying cortical representational remodeling. The objective of the present review is to put various facets of cortical map plasticity into perspective so as to examine possible links between changes occurring at multiple scales of the neural organization of the mature brain. The main focus is on neural substrates that mediate the instructive influence of experience and behavioral context on cortical reorganization, and perceptual correlates of representational remodeling.

Abstract: Contiguous skin surfaces that tend to be synchronously stimulated are represented in neighbouring sectors of primary somatosensory maps. Moreover, neuronal receptive fields (RFs) are reshaped through ongoing competitive/cooperative interactions that segregate/desegregate inputs converging onto cortical neuronal targets. The present study was designed to evaluate the influence of spatio-temporal constraints on somatotopic map organization. A vascularized and innervated pedicle flap of the ventrum skin bearing nipples was rotated by 180 degrees . Electrophysiological maps of ventrum skin were elaborated in the same rats at 24 h after surgery and 2 weeks after parturition. Neurones with split RFs resulting from the surgical separation of formerly adjoining skin surfaces were more numerous in non-nursing than nursing rats. RFs that included newly adjacent skin surfaces on both sides of the scar line emerged in nursing rats, suggesting that the spatial contiguity of formerly separated skin surfaces induced a fusion of their cortical representations through nursing-induced stimulation. In addition, nursing-dependent inputs were found to reincorporate the rotated skin flap representation in an updated topographical organization of the cortical map. A skin territory including recipient and translocated skin areas was costimulated for 7 h, using a brushing device. Neural responses evoked by a piezoelectric-induced skin indentation before and after skin brushing confirmed the emergence of RFs crossing the scar line and contraction of non-brushed components of split RFs. Our findings provide further evidence that the spatiotemporal structure of sensory inputs changing rapidly or evolving in a natural context is critical for experience-dependent reorganization of cortical map topography.

Abstract: We describe a novel stimulus delivery system designed to present tactile stimuli to a subject in the tunnel of a magnetic resonance imaging (MRI) system. Using energy from an air-driven piston to turn a wheel, the device advances a conveyor belt with a pre-determined sequence of stimuli that differ in their spatial features into the tunnel of the MRI. The positioning of one or several stimulus objects in a window near the subject's hand is controlled by a photoelectric device that detects periodic openings in the conveyor belt. Using this electric signal to position each presentation avoids cumulative positioning errors and provides a signal related to the progression of the experiment. We used a series of shapes that differed in their spatial features but the device could carry stimuli with a diversity of shapes and textures. This flexibility allows the experimenter to design a wide variety of psychophysical experiments in the haptic world and possibly to compare and contrast these stimuli with the cognitive treatment of similar stimuli delivered to the other senses. Appropriate experimental design allows separation of motor, sensory and memory storage phases of mental processes.

Abstract: This study is an attempt to gain insight into the malleability of representational maps in the primary somatosensory cortex in relation to the expression of proteins involved in inhibitory and excitatory neurotransmitter systems that contribute to maintain these maps in a dynamic state. Malleability of somatosensory maps is characterized by changes in the sizes of neuron receptive fields (RFs) affecting the representational grain and in the locations and submodalities of these RFs modifying the map extent. The concomitance of these alterations remains so far hypothetical. We used nursing as an evolving source of ethologically significant cutaneous stimulation. This cyclic behavior is particularly suited to investigating the time course of experience-dependent cortical changes. Electrophysiological maps of the ventrum skin were recorded twice in the same lactating rats between nursing initiation and several weeks after nursing. We found that reduction in RF size occurred earlier than map expansion. As nursing time declined, the map expansion was maintained longer than the RF sharpening. Based on this difference in time course, we compared the expression patterns of several activity-dependent proteins in relation to the RF plasticity. Western blot analysis showed an increase in glutamic acid decarboxylase expression that was concomitant with RF contraction. In contrast, NR2A subunit of NMDA and alpha calcium/calmodulin kinase type II were upregulated at times when map expansion was observed. We propose that inhibitory and excitatory plasticity mechanisms operating with different time courses may contribute to the temporal dissociation of nursing-induced RF reshaping and map expansion.

Abstract: We combined behavioral assessment of texture discrimination and electrophysiological mapping of concomitant reorganization in the forepaw representation within the SI cortex. Rats were housed in enriched (EE) or impoverished (IE) environments which have been shown to remodel the forepaw map and possibly alter discriminative abilities. In addition, animals were trained to discriminate homogeneous floorboards of invariant roughness from heterogeneous floorboards of gradually decreasing roughness contrasts during locomotion. As reported recently, differences in perceptual abilities were not related to housing conditions, but to a predilection for a floorboard type [Bourgeon S, Xerri C, Coq JO. Abilities in tactile discrimination of textures in adult rats exposed to enriched or impoverished environments. Behav Brain Res 2004;153:217-231]. Consistently, the present study shows that cortical map remodeling resulting from short-duration daily experience can prevail over changes induced by housing conditions. The relative area of glabrous skin representation was related to the discrimination performance and learning abilities in the rats (H) with a predilection for heterogeneous floorboards, i.e. in the animals performing discrimination in the most challenging perceptual context. By contrast, this cortical area was influenced by the duration of sensory experience in rats (h) with a predilection for homogeneous floorboards. Both EE condition and training to discrimination selectively decreased the sizes of the SI neurons' receptive fields (RFs) located on glabrous skin. Smaller RFs and larger cortical areas serving glabrous skin were correlated with better perceptual performances and learning abilities in the H rats only. The present study shows that representational reorganization related to tactile discrimination performances depends upon the perceptual context.

Abstract: In previous studies, we have shown that housing in enriched environment for about 3 months after weaning improved the topographic organization and decreased the size of the receptive fields (RFs) located on the glabrous skin surfaces in the forepaw maps of the primary somatosensory cortex (SI) in rats [Exp. Brain Res. 121 (1998) 191]. In contrast, housing in impoverished environment induced a degradation of the SI forepaw representation, characterized by topographic disruptions, a reduction of the cutaneous forepaw area and an enlargement of the glabrous RFs [Exp. Brain Res. 129 (1999) 518]. Based on these two studies, we postulated that these representational alterations could underlie changes in haptic perception. Therefore, the present study was aimed at determining the influence of housing conditions on the rat's abilities in tactile texture discrimination. After a 2-month exposure to enriched or impoverished environments, rats were trained to perform a discrimination task during locomotion on floorboards of different roughness. At the end of every daily behavioral session, rats were replaced in their respective housing environment. Rats had to discriminate homogeneous (low roughness) from heterogeneous floorboards (combination of two different roughness levels). To determine the maximum performance in texture discrimination, the roughness contrast of the heterogeneous texture was gradually reduced, so that homogeneous and heterogeneous floorboards became harder to differentiate. We found that the enriched rats learned the first steps of the behavioral task faster than the impoverished rats, whereas both groups exhibited similar performances in texture discrimination. An individual "predilection" for either homogeneous or heterogeneous floorboards, presumably reflecting a behavioral strategy, seemed to account for the absence of differences in haptic discrimination between groups. The sensory experience depending on the rewarded texture discrimination task seems to have a greater influence on individual texture discrimination abilities than the sensorimotor experience related to housing conditions.

Abstract: We used a kinematic analysis for assessing locomotor impairments and evaluating the time course of recovery after focal injury to the forepaw area of the primary somatosensory cortex (SI) in rats. The animals were trained to traverse a beam that was rotated at various speeds. Changes in orientation of the body and independent movement of the anterior and posterior parts of the body were reconstructed using a 3D motion analysis. In addition, we used electrophysiological cortical mapping to search for neurophysiological changes within the spared cortical zones surrounding the lesion. Neuronal recordings were performed in the same animals prior to and 3 weeks after the lesion induction. Our findings show that a focal lesion that destroyed about 60% of the forepaw representational zone was sufficient to cause conspicuous impairments in the rats' ability to produce adequate motor adjustments to compensate for the lateral shift of the beam and to avoid falling. The main deficits were reflected in a lack of appropriate coordination between the anterior and posterior parts of the body and an inability to maintain a regular gait during locomotion. Skilled locomotion was fully recovered within a 2-3 week period. Functional recovery cannot be ascribed to a restitution of the lost sensory representations. A permanent decrease of forepaw representation was recorded despite the re-emergence of restricted representational sectors in the peri-lesion zone. We suggest that alterations may have occurred in other cortical and subcortical areas interconnected with the injured area. It is also conceivable that the functional recovery involved an increased reliance on all the available sources of sensorimotor regulation as well as the use of behavioral strategies.

Abstract: The influence of housing in an enriched or impoverished environment and anti-ischemic treatment (piracetam) on the organization of the intact regions of the somatosensory cortical maps adjacent to a focal cortical injury were investigated in adult rats. Response properties of small clusters of neurons were recorded in the area of the primary somatosensory cortex (SI) devoted to the contralateral forepaw representation. Electrophysiological maps were elaborated on the basis of the sensory "submodality" (cutaneous or noncutaneous) and the location of the receptive fields (RFs) of layer IV neurons. Recordings were made before, and 3 weeks after induction of a focal neurovascular lesion to the SI cortex. The main results were: 1) the focal ischemic injury induced a cellular loss which was less severe in the piracetam treated rats, regardless of the housing conditions; 2) the lesion resulted in a compression of the remaining forepaw map, a fragmentation of the representational zones serving the cutaneous surfaces (low-threshold inputs) and an enlargement of noncutaneous zones (high-threshold inputs) in the spared cortical sectors surrounding the lesion. These changes were found in all placebo rats, with the most detrimental effects in the animals exposed to an impoverished environment, and in the piracetam-plus-impoverished rats. In contrast, a limited compression of the forepaw map and a preservation of most representational sectors were observed in the piracetam-plus-enriched animals, 3) the size of the cutaneous RFs of the neurons within the intact cortical zones remained unchanged, regardless of environment or treatment; 4) consistent with the map changes, the skin surfaces lacking low-threshold cutaneous RFs increased after the lesion in all animal groups but the piracetam-plus-enriched rats; 5) cortical responsiveness as assessed with mechanical threshold evaluation was diminished in the placebo rats, whatever the environment, and in the piracetam-impoverished rats, but was not significantly affected in the piracetam-enriched animals. Our findings, based on the first double electrophysiological mapping in the rat SI cortex, highlight the protective effects of an environmental therapy associated with an anti-ischemic treatment on the neurophysiological properties of cortical neurons following a focal neurovascular injury to the neocortex.

Abstract: Acute and chronic postlesion reorganization of the cortical maps was examined in adult rats using electrophysiological mapping of the forepaw area in the primary somatosensory cortex. Recordings were made before and after (first 12 hr and 3 wk) induction of a focal thermal-ischemic lesion to a restricted part of the forepaw area. The influence of an anti-ischemic substance (piracetam) and housing in an enriched environment (EE) or impoverished environment (IE) on the organization of the spared regions of the cortical maps adjacent to the lesion was investigated. The results revealed (1) a gradual expansion of the injured zone and a cellular loss that were smaller in the piracetam-treated (PT) rats than in the placebo (PL) rats; (2) a better preservation of the somatotopic organization and the neuronal responsiveness in the maps of the PT rats during the first 12 hr after the lesion; (3) a gradual compression and fragmentation of the remaining forepaw map over the first 3 postlesion wk. These changes were found in all PL rats, with the most detrimental effects in the animals exposed to an IE. In the PT-EE animals, a contrasting substantial preservation of the map was observed. (4) Cortical responsiveness was diminished in the PL rats, whatever the environment, and in the PT-IE rats; but it was not significantly affected in the PT-EE animals. The findings demonstrate the protective function of acute piracetam treatment on electrophysiological properties of cortical neurons within the peri-infarct tissue and highlight the neuroprotective effects of an environmental therapy combined with the piracetam treatment during the weeks after ischemic damage.

Abstract: In a previous study, we found that the forepaw representation in the primary somatosensory cortex of rats housed in standard laboratory conditions was drastically altered during the aging process. In other studies we reported that exposure to an enriched environment improved the topographical organization and increased the spatial resolution of the forepaw cutaneous map in young adult rats, whereas housing in impoverished environment resulted in a loss of somatotopic details in the forepaw map. The main purpose of the present study was to investigate the influence of differential sensorimotor experience promoted by exposure to enriched or impoverished environments on the mutability of the cortical forepaw representation during aging. Two groups of Long-Evans rats were reared in enriched and impoverished environments from weaning to the age of 3.5-5 months (young adults), 6.5-8 months (mature rats), and 23-28 months (senescent rats). The electrophysiological maps of the forepaw representation were based on the somatosensory 'submodality' (cutaneous vs. non-cutaneous), size, and location of the receptive fields of small clusters of layer IV neurons. Moreover, the mechanical thresholds of neuronal response to cutaneous stimulation were assessed with calibrated von Frey filaments in mature and senescent animals. Age-related alterations of the topographic features of the forepaw map were characterized by a decrease in and a fragmentation of the cortical zones serving the glabrous skin of the forepaw. These changes were less pronounced in the enriched rats than in the impoverished rats. Glabrous skin receptive fields were smaller in young adult and mature enriched rats than in their impoverished counterparts. However, during aging glabrous receptive fields increased in the enriched rats, but decreased in the senescent impoverished rats so that old rats of either groups displayed receptive fields of similar sizes; in contrast, the size of hairy skin receptive fields was not affected by housing conditions or aging. Measurement of the neuronal responses to calibrated forces applied to the skin indicated that cortical excitability to near-threshold cutaneous input was lower in senescent rats than in mature rats, regardless of environmental conditions.The present study demonstrates that use-dependent remodeling of somatosensory maps occurs throughout life and that environmental and social interactions can partially offset the age-related breakdown of somatosensory cortical maps.

Abstract: The cortical forepaw representation of adult rats was mapped by using multiunit recordings of layer IV neurons within the primary somatosensory cortex. Electrophysiological maps were based on somatosensory "submodality" (cutaneous vs non-cutaneous), location and size of the receptive fields. Age-related changes in the organizational features of the forepaw representation in the primary somatosensory cortex were analysed in adult rats whose ages ranged from 3.5 to five months (young), about eight months (mature), 15-17months (presenescent) to 24-28months (senescent). Rats were housed from weaning (30days postnatal) in a standard laboratory environment. The organization of the forepaw map was not gradually altered with advancing age, but striking changes occurred in early adulthood (before eight months) and did not progress with further aging. The main alterations consisted of a prominent decrease in, and a fragmentation of, the cutaneous area of the forepaw representation. Representational zones formerly serving cutaneous surfaces became predominantly activated by high-threshold, presumably non-cutaneous, inputs which appeared somatotopically organized. These emergent non-cutaneous zones were interspersed with cutaneous sectors, thereby disrupting the somatotopic organization of the map of the forepaw skin. No significant modification in the size of glabrous or hairy cutaneous receptive fields accompanied these changes. Subjective evaluation of the responses evoked by tactile stimulation suggests that neuronal responsiveness was increased in the eight- to 17-month-old rats, but less so in the 24- to 28-month-old animals.These results indicate that degradation of the somatotopic organization of the cutaneous representation of the forepaw in the rat somatosensory cortex occurs early during the course of adult life.

Abstract: We investigated the effects of sensory deprivation on the forepaw representation in the primary somatosensory cortex (SI) in the adult rat. Cortical maps were constructed from high-resolution multiunit recordings of the response of layer IV neurons to somatosensory stimuli. The main features of the forepaw representation were described in terms of areal extent and topography of the cortical map, and sensory submodality, size, and location of the receptive field (RF) of small clusters of the cortical neurons. After being weaned, two groups of Long-Evans rats were housed in a standard (SE) or impoverished (IE) environment for 65-115 days. A third group of SE rats was subjected to severe sensorimotor restriction (SR) of one forepaw for 7 days or 14 days, by using a one-sleeved cast. A concomitant effect of unilateral forelimb immobilization was a forced use of the nonrestricted forelimb in postural balance. The maps of both forepaws were derived 24 h after the cast was removed and the animal was allowed normal limb use. In a fourth group, SE rats experienced a 7-day immobilization followed by symmetrical limb use for 7 days before we mapped the hemisphere contralateral to the casted limb. For the SE and IE rats, the total areal extent of the cutaneous forepaw representation was similar, but IE rats exhibited a significant expansion of cortical islets serving high-threshold, presumably noncutaneous inputs, which were included in the cutaneous maps. In addition, SI neurons of IE rats had greatly enlarged glabrous, but not hairy, skin RFs. For the SR rats, the areal extent of the cutaneous map of the casted forepaw decreased by about 50%, after both 7- and 14-day forelimb immobilization. Large cortical sectors presumed to be formerly activated by cutaneous inputs were driven by high-threshold inputs that disrupted the somatotopic representation of the forepaw skin surfaces. These "emergent" representational sectors were topographically organized. By contrast, the areal extent and topography of the non-casted forepaw representation did not differ from those of SE rats. The size of glabrous RFs on the casted forepaw was similar to that of SE rats. On the contrary, glabrous RFs on the noncasted forepaw of SR rats were larger than those on their casted forepaw. The size of hairy RFs was not altered by the forelimb restriction. Interestingly, alteration of the somatotopic features of the casted forepaw map persisted after 7 days of symmetric use of the forelimbs. The present study demonstrates that continuous sensory experience is needed for the organizational features of SI maps to be maintained.

Abstract: The representations of the surfaces of the hand in the primary somatosensory cortical field, area 3b, were reconstructed in detail in seven owl monkeys and two squirrel monkeys trained to pick up food pellets from five wells of different sizes. From an early clumsy performance in which several to many retrieval attempts were required for each successful pellet retrieval, the monkeys exhibited a gradual improvement in digital dexterity as shown by significant decreases in mean numbers of grasp attempts/successful retrieval and corresponding standard deviations (e.g. 5.8 +/- 4.5 and 4.8 +/- 3.1 respectively, for the smallest well) between the first and last training sessions. All monkeys commonly used alternative, specific retrieval strategies involving various combinations of digits for significant time epochs before developing a highly successful strategy, which, once achieved, was rapidly stereotyped. For example, the numbers of digit combinations used during the first five versus the last five training sessions decreased from 3.3 +/- 0.7 to 1.8 +/- 0.6 for the smallest well. In both owl and squirrel monkeys, as the behavior came to be stereotyped, monkeys reliably engaged limited surfaces of the glabrous tips of two digits (in eight monkeys), or of three digits (in one monkey) in the palpation and manipulation of these small pellets for their location, capture, and transportation to the mouth. In cortical area 3b, the magnification of representation of these differentially engaged glabrous fingertip surfaces was nearly 2x larger than for the corresponding surfaces of other hand digits, or for the contralateral cortical representations of the same digit surfaces on the opposite hand. In parallel, cutaneous receptive field for area 3b neurons representing crucial digital tip surfaces were less than half as large as were those representing the corresponding surfaces of control digits. Receptive field overlaps were smaller on the trained fingertips than on control fingers. Moreover, the proportion of small overlaps was greater for the trained digits (76 +/- 7%) than for the other digits of the same hand (49 +/- 5.4%). There was still a simple, single--but apparently topologically expanded--representation of these differentially engaged skin surfaces in these monkeys. Thus, with very limited manual exercise over a total period of a few hours of practice at a skill played out in brief daily sessions over a several week long training period, the representations of skin surfaces providing information crucial for successfully performing a small-object retrieval behavior appeared to be substantially remodeled in the most 'primary' of the SI somatosensory cortical fields, cortical area 3b. By that remodeling, behaviorally important skin surfaces were represented in a much finer representational grain than normal. Some implications of these findings for motor skill acquisition are discussed.

Abstract: Immediate postlesion reorganization of the somatosensory cortical representation was examined in adult rats. Response properties of small clusters of neurons were recorded in the area of the primary somatosensory cortex (SI) devoted to the contralateral forepaw representation. Electrophysiological maps were elaborated on the basis of the sensory 'submodality' (cutaneous or noncutaneous) and the location of the peripheral receptive fields (RFs) of layer IV neurons. Recordings were made prior to, and from 1 to 12 h after, induction of a focal neurovascular lesion to the SI cortex that initially destroyed a part (8.5%) of the cutaneous representation. Moreover, the influence of an anti-ischaemic substance (piracetam) on lesion-induced changes was analysed. The main observations were: (i) a gradual outward expansion of the area of the functional lesion, which was smaller in the piracetam-treated (PT) rats than in the control, placebo-treated (PL) rats; (ii) a substantial remodelling of the spared representational zones, both in cortical sectors adjoining the site of injury and those remote from the site; (iii) a significant postlesion increase in the size of cutaneous RFs in the PT rats, but not in the PL rats; (iv) a better preservation of RF submodality and topographic organization in the PT maps than in the PL maps; and (v) a decrease in neuronal responsiveness to cutaneous stimulation which was less pronounced in the PT than in the PL rats. Our results can be ascribed to a rapid change in the balance of excitatory and inhibitory connections which leads to unmasking of subthreshold inputs converging onto cortical neurons. Our findings also indicate that acute piracetam treatment exerts a protective function on the physiological response properties of cortical neurons after focal injury.

Abstract: Adult owl and squirrel monkeys were trained to master a small-object retrieval sensorimotor skill. Behavioral observations along with positive changes in the cortical area 3b representations of specific skin surfaces implicated specific glabrous finger inputs as important contributors to skill acquisition. The area 3b zones over which behaviorally important surfaces were represented were destroyed by microlesions, which resulted in a degradation of movements that had been developed in the earlier skill acquisition. Monkeys were then retrained at the same behavioral task. They could initially perform it reasonably well using the stereotyped movements that they had learned in prelesion training, although they acted as if key finger surfaces were insensate. However, monkeys soon initiated alternative strategies for small object retrieval that resulted in a performance drop. Over several- to many-week-long period, monkeys again used the fingers for object retrieval that had been used successfully before the lesion, and reacquired the sensorimotor skill. Detailed maps of the representations of the hands in SI somatosensory cortical fields 3b, 3a, and 1 were derived after postlesion functional recovery. Control maps were derived in the same hemispheres before lesions, and in opposite hemispheres. Among other findings, these studies revealed the following 1) there was a postlesion reemergence of the representation of the fingertips engaged in the behavior in novel locations in area 3b in two of five monkeys and a less substantial change in the representation of the hand in the intact parts of area 3b in three of five monkeys. 2) There was a striking emergence of a new representation of the cutaneous fingertips in area 3a in four of five monkeys, predominantly within zones that had formerly been excited only by proprioceptive inputs. This new cutaneous fingertip representation disproportionately represented behaviorally crucial fingertips. 3) There was an approximately two times enlargement of the representation of the fingers recorded in cortical area 1 in postlesion monkeys. The specific finger surfaces employed in small-object retrieval were differentially enlarged in representation. 4) Multiple-digit receptive fields were recorded at a majority of emergent, cutaneous area 3a sites in all monkeys and at a substantial number of area 1 sites in three of five postlesion monkeys. Such fields were uncommon in area 1 in control maps. 5) Single receptive fields and the component fields of multiple-digit fields in postlesion representations were within normal receptive field size ranges. 6) No significant changes were recorded in the SI hand representations in the opposite (untrained, intact) control hemisphere. These findings are consistent with "substitution" and "vicariation" (adaptive plasticity) models of recovery from brain damage and stroke.

Abstract:

Abstract: The cortical forepaw area of young adult rats was mapped by recording the response properties of small clusters of neurons in layer IV of the primary somatosensory (SI) cortex. First we quantitatively analyzed the somatotopic organizational features of the cortical forepaw representation in terms of areal extent and topography, receptive field (RF) sensory modality, size, and location. We also assessed the influence of environmental enrichment, known to induce structural alterations in cortical connectivity, on the representational characteristics of the forepaw maps. Long-Evans rats were housed in environments (standard, SE; enriched, EE) promoting differential tactile experience for 71-113 days from weaning. Within the SI, we found a single and complete topographic map of the cutaneous surfaces of the forepaw consisting of a rostrolateral-caudomedial sequence of digit and pad representational zones. Small islets of noncutaneous responses (NCR; high-threshold, deep-receptor input) within the boundaries of the cutaneous maps were a conspicuous feature of the forepaw map for SE rats. These islets created discontinuities in the representation of contiguous skin territories. In the SE rats, about 79% of the cortical sites activated by light tactile stimulation had a single cutaneous RF, whereas about 21% exhibited multiple RFs. Most single-digit RFs we delineated in the SE rats extended across two or three phalanges. As a result, the representations of the phalangeal skin surfaces were not segregated but formed an overlapping continuum. Moreover, within these regions, as the electrode was displaced in regular steps across the mediolateral axis, RFs did not shift across the digit skin surface in an orderly manner, suggesting a lack of internal topography in the finger representation zones. Tactile experience promoted by environmental enrichment induced alterations in the representational features of the SI cutaneous map of the forepaw. In EE rats, the areal extent of the forepaw cutaneous representation was 1.5 times larger than in SE rats. Indeed, the cutaneous map extended into NCR cortical sectors along its external margins and also into NCR islets found in the forepaw area. Consequently, in EE rats there were fewer representational discontinuities. The areal enlargement was due to a selective increase in the areal extent of the glabrous but not the hairy skin surface representations. Furthermore, protuberant glabrous skin (digit tips, palmar pads) was represented over larger cortical regions than were other glabrous skin territories less likely to be stimulated during object palpation and manipulation. Maps from EE rats were also characterized by a larger proportion of sites with single RFs (88% compared with 79%). In addition, glabrous RFs from EE rats were smaller and more clustered on the digit tips and palmar pads than were RFs in SE rat maps. RF size on hairy skin surfaces remained unchanged. Because the RFs were smaller, the cutaneous maps of EE rats contained distinct representations of digit phalangeal glabrous skin. RFs tended to exhibit more orderliness in their progression across the digit glabrous skin of EE rats than they did in SE rats. The phalanges of EE rats were represented in distinct patches. Neurons in EE rats were more sensitive to light tactile stimulation than were neurons in SE rats. These alterations were presumably mediated by the selective potentiation of cutaneous over deep-receptor activation. More generally, the present study corroborates the view that cortical cutaneous maps are maintained in a permanent state of use-dependent fluctuation.

Abstract: Vestibular inputs tonically activate the anti-gravitative leg muscles during normal standing in humans, and visual information and proprioceptive inputs from the legs are very sensitive sensory loops for body sway control. This study investigated the postural control in a homogeneous population of 50 unilateral vestibular-deficient patients (Ménière's disease patients). It analyzed the postural deficits of the patients before and after surgical treatment (unilateral vestibular neurotomy) of their diseases and it focused on the visual contribution to the fine regulation of body sway. Static posturographic recordings on a stable force-plate were done with patients with eyes open (EO) and eyes closed (EC). Body sway and visual stabilization of posture were evaluated by computing sway area with and without vision and by calculating the percentage difference of sway between EC and EO conditions. Ménière's patients were examined when asymptomatic, 1 day before unilateral vestibular neurotomy, and during the time-course of recovery (1 week, 2 weeks, 1 month, 3 months, and 1 year). Data from the patients were compared with those recorded in 26 healthy, age- and sex-matched participants. Patients before neurotomy exhibited significantly greater sway area than controls with both EO (+52%) and EC (+93%). Healthy participants and Ménière's patients, however, displayed two different behaviors with EC. In both populations, 54% of the subjects significantly increased their body sway upon eye closure, whereas 46% exhibited no change or significantly swayed less without vision. This was statistically confirmed by the cluster analysis, which clearly split the controls and the patients into two well-identified subgroups, relying heavily on vision (visual strategy, V) or not (non-visual strategy, NV). The percentage difference of sway averaged +36.7%+/-10.9% and -6.2%+/-16.5% for the V and NV controls, respectively; +45.9%+/-16.8% and -4.2%+/-14.9% for the V and NV patients, respectively. These two distinct V and NV strategies seemed consistent over time in individual subjects. Body sway area was strongly increased in all patients with EO early after neurotomy (1 and 2 weeks) and regained preoperative values later on. In contrast, sway area as well as the percentage difference of sway were differently modified in the two subgroups of patients with EC during the early stage of recovery. The NV patients swayed more, whereas the V patients swayed less without vision. This surprising finding, indicating that patients switched strategies with respect to their preoperative behavior, was consistently observed in 45 out of the 50 Ménière's patients during the whole postoperative period, up to 1 year. We concluded that there is a differential weighting of visual inputs for the fine regulation of posture in both healthy participants and Ménière's patients before surgical treatment. This differential weighting was correlated neither with age or sex factors, nor with the clinical variables at our disposal in the patients. It can be accounted for by a different selection of sensory orientation references depending on the personal experience of the subjects, leading to a more or less heavy dependence on vision. The change of sensory strategy in the patients who had undergone neurotomy might reflect a reweighting of the visual and somatosensory cues controlling balance. Switching strategy by means of a new sensory selection of orientation references may be a fast adaptive response to the lesion-induced postural instability.

Abstract: In a first study, the representations of skin surfaces of the hand in the primary somatosensory cortex, area 3b, were reconstructed in owl monkeys and squirrel monkeys trained to pick up food pellets from small, shallow wells, a task which required skilled use of the digits. Training sessions included limited manual exercise over a total period of a few hours of practice. From an early clumsy performance in which many retrieval attempts were required for each successful pellet retrieval, the monkeys exhibited a gradual improvement. Typically, the animals used various combinations of digits before developing a successful retrieval strategy. As the behavior came to be stereotyped, monkeys consistently engaged surfaces of the distal phalanges of one or two digits in the palpation and capture of food pellets from the smallest wells. Microelectrode mapping of the hand surfaces revealed that the glabrous skin of the fingertips predominantly involved in the dexterity task was represented over topographically expanded cortical sectors. Furthermore, cutaneous receptive fields which covered the most frequently stimulated digital tip surfaces were less than half as large as were those representing the corresponding surfaces of control digits. In a second series of experiments, Long-Evans rats were assigned to environments promoting differential tactile experience (standard, enriched, and impoverished) for 80 to 115 days from the time of weaning. A fourth group of young adult rat experienced a severe restriction of forepaw exploratory movement for either 7 or 15 days. Cortical maps derived in the primary somatosensory cortex showed that environmental enrichment induced a substantial enlargement of the cutaneous forepaw representation, and improved its spatial resolution (smaller glabrous receptive fields). In contrast, tactile impoverishment resulted in a degradation of the forepaw representation that was characterized by larger cutaneous receptive fields and the emergence of non-cutaneous responses. Cortical maps derived in the hemispheres contralateral to the immobilized forelimb exhibited a severe decrease of about 50% in the overall areal extent of the cutaneous representation of the forepaw, which resulted from the invasion of topographically organized cortical zones of non-cutaneous responses, and numerous discontinuities in the representation of contiguous skin territories. The size and the spatial arrangement of the cutaneous receptive fields were not significantly modified by the immobilization of the contralateral forelimb. Similar results were obtained regardless of whether the forelimb restriction lasted 7 or 15 days. These two studies corroborate the view that representational constructs are permanently reshaped by novel experiences through dynamic competitive processes. These studies also support the notion that subject-environment interactions play a crucial role in the maintenance of basic organizational features of somatosensory representations.

Abstract: Vision has long been recognized as a sensorimotor system which plays a major role in substitution for functional deficits induced by unilateral or bilateral exclusion of primary vestibular afferents. Little is known, however, about the post-lesion influence of visual inputs on the recovery of posturo-kinetic balance in a situation where fine, well-coordinated locomotor adjustments are required. The present study was carried out in order to gain some insight into the role played by motion vision in the restoration of fine posturo-kinetic balance in adult cats subjected to unilateral vestibular neurectomy. Prior to the lesion, 15 adult animals were trained to cross a beam rotating at various speeds. Their best global balance performance (highest beam rotation speed that did not provoke falling) and their average locomotion speed were evaluated. After the lesion, the cats were separated into three groups: (1) five animals were placed in a normal environment (animal house) (NV cats); (2) four animals were exposed to stroboscopic illumination which eliminated visual motion cues (SV cats) for 2 weeks following the lesion; and (3) three animals were placed in a normal environment and their training was interrupted for the same period as in the SV cats (NVI cats). The possible influence of the 2-week deprivation of visual motion cues on posturo-kinetic balance was also examined in three intact cats. The present behavioral study showed that: (1) early sensory deprivation caused suspension of the posturo-kinetic balance recovery process as long as it was maintained; (2) complete restoration of global balance capacities developed following the vestibular neurectomy after a significant delay in half of the SV cats; (3) the lack of motion cues resulted in severe alterations of fine posturo-kinetic balance (inappropriate dynamic motor adjustments and irregular locomotion speed regulation) in all SV cats; and (4) the visual deprivation induced a 2-week delay in the restoration of fine locomotor balance. These findings provide evidence for a defect in the visual sensory substitution processes that normally take place within the first few weeks following exclusion of primary vestibular afferents.

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Abstract: The purpose of this study was to investigate adaptive changes in the activity of vestibular nuclei neurons unilaterally deprived of their primary afferent inputs when influenced by visual motion cues. These neuronal changes might account for the established role that vision plays in the compensation for posturo-kinetic deficits after the loss of vestibular inputs. Neuronal recordings were made in alert, non-paralysed cats that had undergone unilateral vestibular nerve sections. The unit responses collected in both Deiters' nuclei were compared to those previously recorded in intact cats. We analysed the extracellular activity of Deiters' nucleus neurons, as well as the optokinetic reflex (OKR) evoked during sinusoidal translation of a whole-field optokinetic stimulus in the vertical plane. In intact cats, we found the unit firing rate closely correlated with the visual surround translation velocity, and the relationship between the discharge rate and the motion frequency was tuned around an optimal frequency. The maximum firing rate modulation was generally below the 0.25 Hz stimulus frequency; unit responses were weak or even absent above 0.25 Hz. From the 4th day to the end of the 3rd week after ipsilateral deafferentation, a majority of cells was found to display maximum discharge modulation during vertical visual stimulation at 0.50 Hz, and even at 0.75 Hz, indicating that the frequency bandwidth of the visually induced responses of deafferented vestibular nuclei neurons had been extended. Consequently, the frequency-dependent attenuation in the sensitivity of vestibular neurons to visual inputs was much less pronounced. After the first 3 weeks post-lesion, the unit response characteristics were very similar to those observed prior to the deafferentation. On the nucleus contralateral to the neurectomy, the maximum modulation of most cells was tuned to the low frequencies of optokinetic stimulation, as also seen prior to the lesion. We found, however, a subgroup of cells displaying well-developed responses above 0.50 Hz. Under all experimental conditions, the neuronal response phase still remained closely correlated with the motion velocity of the vertical sinusoidal visual pattern. We hypothesize that Deiters' neurons deprived of their primary afferents may transiently acquire the ability to code fast head movements on the basis of visual messages, thus compensating, at least partially, for the loss of dynamic vestibular inputs during the early stages of the recovery process.(ABSTRACT TRUNCATED AT 400 WORDS)

Abstract: The representation of the surfaces of the trunk was mapped in detail in a series of anesthetized adult female rats to assess cortical representational changes that might be induced in the SI cortical field by a major natural source of a differentially heavy schedule of tactile inputs: the stimulation of the rat ventrum in nursing behavior. Controls included virgin rats and postpartum age-matched rats whose litters were removed on the day of birth. The SI representation of the ventral trunk skin of lactating rats was about 1.6 x larger than in matched postpartum nonlactating or virgin controls. The greatest representational change--about twofold--was for the nipple-bearing skin between the forelimbs and hindlimbs. Indeed, changes in SI representational territory for the middle third of the ventrum, a skin zone without nipples, were not significant. As a rule, the representation of the ventrum skin in lactating rats was at least as topographically ordered as was that reconstructed for nonlactating postpartum and virgin controls. Receptive fields (RFs) representing the ventrum skin in lactating females were about one-third the sizes of those recorded in matched nonlactating or virgin controls. RF size differences were again greater for the representation of the nipple-bearing skin in the anterior and posterior thirds of the ventrum than for the central third. Changes in RF sizes were roughly inversely related to changes in the cortical magnification of representation of the ventrum on the proportion of about 3:2. Interestingly, the glabrous nipple and areolar skin were only weakly represented--or not demonstrably represented--in the SI map of either lactating or control rats. These results indicate that there is a largely unstudied cortical neurology of nursing behavior. Major CNS changes are induced by this dramatic, episodic change in behaviorally important tactile inputs. In turn, input-induced changes presumably contribute to this emergent, rapidly evolving behavior.

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Abstract: Extracellular activity from vestibular nuclei neurons and vertical eye movements were recorded in the alert cat during sinusoidal optokinetic stimulation in the vertical plane at frequencies varying from 0.0125 Hz to 0.75 Hz. Among a population of 96 vestibular units located in and around Deiters' nucleus, 73 neurons (76%) displayed a firing rate modulation which followed the input at the standard parameters of visual stimulation (0.05 Hz; 10.1 deg/s or 9.1 cm/s peak to peak velocity). Two different patterns of modulation were found. In 42 cells (57%) an increase in the firing rate was observed during motion of the visual scene in the downward direction, while 31 neurons (43%) showed the opposite behavior, with an enhanced firing rate during upward movement. The phase of the neuronal responses was close (+/- 45 degrees) to the velocity peaks (+90 degrees: downward and -90 degrees: upward) of visual scene motion for 65 among the 73 neurons. Mean values of phase was -6.1 +/- 19.5 degrees (SD) and -3.2 +/- 15.5 degrees (SD) with respect to the +90 degrees and -90 degrees velocity peaks, respectively. In the frequency range 0.0125-0.75 Hz, the phase of the neuronal responses remained almost stable, with only a slight lag which reaches -22 degrees at the 0.25 Hz visual stimulation. The firing rate modulation was found to be predominant at low frequencies (0.0125 Hz-0.25 Hz), with three distinct peaks of modulation occurring either at 0.025 Hz, 0.10 Hz or 0.25 Hz, depending on the recorded cells. Above 0.5 Hz, the cell modulation was very poorly developed or even absent. A gain attenuation was observed in all units, which was more important in cells showing a peak of modulation at 0.025 Hz as compared with the others (-20.7 dB vs -9.6 dB, respectively, in the 0.025 Hz-0.25 Hz decade). The gain of the optokinetic reflex (OKR) progressively decreased from mean values of 0.78 +/- 0.15 to 0.05 +/- 0.06 in the 0.025 Hz-0.5 Hz frequency range. A close correlation was observed between the OKR slow phase velocity and the modulation of the neuronal responses in the two cell populations with maximal modulations at 0.10 Hz or 0.25 Hz. No correlations were noticed in the third population characterized by a peak of modulation at 0.025 Hz. In all units, the phase of eye movement velocity and of neuronal responses were both related to the velocity of the visual surround motion. These correlations were also found when varying the amplitude of the visual stimulation at a fixed frequency.(ABSTRACT TRUNCATED AT 400 WORDS)

Abstract: In the present study we have investigated in the awake cat the response dynamics of vestibular nuclei neurons to visual or/and otolith stimulation elicited by vertical linear motion. Of the 53 units tested during sinusoidal motion at 0.05 Hz (9.1 cm/s), 1 (1.9%) was responsive to the otolith input only, 13 (24.5%) were influenced by the visual input only and 23 (43.4%) responded to both modalities. Neurons were excited either during upward or downward animal or visual surround movement. Most units displayed a firing rate modulation very close to motion velocity. All the neurons receiving convergent visual and otolith inputs (0.05 Hz, 9.1 cm/s) exhibited synergistic patterns of response. Motion velocity coding was improved in terms of input-output phase relationship and response sensitivity when visual and otolith signals were combined. Depending on the units, visual-otolith interactions in single neurons could follow a linear or a nonlinear mode of summation. The dynamic characteristics of visual-otolith interactions were examined in the 0.05 Hz-0.50 Hz frequency bandwidth. Visual signals seemed to predominate over otolith signals at low stimulus frequencies (up to 0.25 Hz), while the contrary was found in the higher frequency range of movement (above 0.25 Hz). The effects of visual stabilization (VS: suppression of visual motion cues) was observed in a small sample of units. As a rule, VS induced a reduction in the amplitude of unit response as compared to visual + otolith stimulation, the lower the motion frequency, the more pronounced the attenuation. VS also decreased the amplitude of the otolith-dependent component of response. The possible modes of visual-vestibular interactions in single cells are discussed. The present study supports the hypothesis that visual and vestibular motion cues are weighted according to their internal relevance.

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Abstract: Electromyographic activity of dorsal neck muscles elicited by sinusoidal vertical linear accelerations was studied in alert cats over a wide range of frequencies. Experiments were performed in head-fixed cats and total darkness in order to activate selectively the otolith system. The polyunitary EMG activity was recorded from splenius capitis muscles in normal and labyrinthectomized cats during vertical translations varying from 0.05-1 Hz with a fixed 290 mm peak-to-peak amplitude. The corresponding accelerations ranged from 0.003-1.2 g. In normal cats, the results showed a bilateral and sinusoidal modulation of the EMG activity characterized by two typical EMG patterns depending on the stimulus frequency. In the low-frequency range (0.05-0.25 Hz), the neck muscles responses were composed of a second harmonic (frequency double that of the input signal: H2 responses). The H2 pattern was characterized by an increase in EMG activity during both the upward and downward parts of translation. These two components of the H2 response were closely related to the two peak velocities (+90 degrees and -90 degrees) of the animal motion. Only slight decreases in amplitude and shifts in phase were observed when increasing the frequency. In the higher frequency range (0.25-1 Hz), the neck muscles response was composed of a fundamental frequency corresponding to the input signal (H1 response). The H1 pattern was in phase with the peak of downward acceleration at 0.25 Hz. A phase lag (up to 45 degrees) and a gain attenuation (16.5 dB) were observed when increasing the frequency. The two H1 and H2 EMG patterns were totally absent in bilateral vestibular neurectomized cats. In unilateral vestibular neurectomized cats, a strong drop in gain and phase advance was noted, which mainly affected the H1 pattern. The present results describe some characteristics of otolith-spinal reflexes acting on the head musculature during vertical motion. They are compared with the neuronal responses that we have recorded within the vestibular nuclei complex in the same experimental conditions. The functional role of the vertical otolith-neck reflexes in stabilizing the head in space during many real-life situations is discussed.

Abstract: The aim of the present study was to investigate some aspects of the central processing of otolith information during linear motion. For this purpose, the response characteristics of 69 vestibular nuclei units to sinusoidal otolith stimulation in the vertical Z axis were analysed in the alert cat. Among this population of neurons which responded to a 0.05 Hz, 290 mm translation, 47 units (70%) displayed a firing rate modulation which followed the input frequency (H1 units). The majority of these neurons exhibited an increase in discharge rate during upward displacement, with a response phase close to the motion velocity or slightly leading downward acceleration. The acceleration related units were divided into two groups according to whether they showed clear increases or only a slight change in discharge rate when the stimulus frequency was increased. The former group was characterized by an average -16.3 dB drop in gain (from 43.9 +/- 1.8 dB, S.D. to 27.6 +/- 7 dB, S.D.) within the 0.05 Hz-0.5 Hz frequency range, while the latter group displayed an average -31.2 dB gain attenuation (from 45.1 +/- 1.1 dB, S.D. to 13.9 +/- 0 dB) within the same decade. In contrast to differences in response gain, all the units tested exhibited a relatively stable phase lead of about 20 degrees with respect to downward peak acceleration. Conversely, units whose response was close to motion velocity in the lower frequency range (0.05 Hz-0.10 Hz) displayed a strong phase lead of about 100 degrees when the stimulus frequency was increased (up to 0.50 Hz). These neurons were thus characterized by an acceleration related response in the higher frequency range. At the same time, an average -24.8 dB gain attenuation (from 47.7 +/- 3.4 dB to 22.9 +/- 3.7 dB) was found in the 0.05 Hz-0.5 Hz decade. The remaining 22 neurons (30%) were called H2 units since they displayed a response waveform double that of the input frequency, a response already described during sinusoidal rotation. Unit discharge reached a peak approximately in phase with maximum upward and downward velocity. Asymmetrical change in unit firing rate about the resting discharge level and different dynamic behavior of the upward and downward response components were usually found. These response characteristics suggest that the H2 patterns are centrally constructed and could result from convergence of otolith afferents having opposite polarization vectors.(ABSTRACT TRUNCATED AT 400 WORDS)

Abstract: The response characteristics of neurons located in the lateral vestibular nucleus (LVN) to neck rotation at 0.026 Hz, 10 degrees peak displacement, have been investigated in precollicular decerebrate cats submitted to ipsilateral acute (aVN) or chronic vestibular neurectomy (cVN). On the whole, 105 units were tested after aVN (i.e., during the first postoperative hours) and 132 units after cVN (i.e., after full compensation of the postural and locomotor deficits). The neurons were histologically located either in the rostroventral (rvLVN) or the dorsocaudal part (dcLVN) of Deiters' nucleus, which are known to project mainly to the cervical and the lumbosacral cord, respectively. Moreover, 55 units in the former group and 66 units in the latter group were identified as vestibulospinal neurons projecting to lumbosacral segments of the spinal cord. The responses of these 237 LVN neurons to the neck input were then compared with those of 120 LVN neurons recorded previously in decerebrate cats with intact labyrinths. Whereas 58.3% of the LVN units recorded in control experiments were responsive to neck rotation, 69.5% of the units were affected by this stimulation at the acute stage of the neurectomy and 74.2% at the chronic stage. This increase in responsive units after aVN and cVN with respect to the controls was found exclusively in the dcLVN. The mean discharge rate of the responsive LVN neurons decreased from 40.7 +/- 48.9 (SD) imp/s in control experiments to 22.1 +/- 15.8 (SD) imp/s after a VN. Similar value was also obtained after cVN [25.0 +/- 17.2 (SD) imp/s], suggesting that compensation of the postural deficits elicited by the vestibular neurectomy results from a redistribution of the excitatory drive within different populations of LVN neurons. Indeed, the relation found in control experiments, i.e., that the faster the conduction velocity of vestibulospinal axons the lower was the unit discharge at rest, was lost after aVN, due to a decrease in resting discharge of the slow units. The mean discharge rate of the slow units, however, recovered after cVN, so that the negative correlation between resting discharge rate and axonal conduction velocity was reestablished. The average gain and sensitivity of the first harmonic response of the LVN neurons to neck rotation recorded after aVN and cVN were comparable to those obtained in preparations with the vestibular nerves intact.(ABSTRACT TRUNCATED AT 400 WORDS)

Abstract: The responses of lateral vestibular nucleus (LVN) neurons to stimulation of macular labyrinth receptors have been investigated in precollicular decerebrate cats after contralateral acute vestibular neurectomy (aVN). On the whole, 78 LVN neurons were tested during slow sinusoidal tilt of the animal at the standard parameters (0.026 Hz, 10 degrees peak displacement). The neurons were located in both the rostroventral (rvLVN) and the dorsocaudal parts (dcLVN) of Deiters' nucleus, which project mainly to the cervical and the lumbosacral cord, respectively. After contralateral aVN, the proportions of responsive units in rvLVN and dcLVN (100% and 75.4%, respectively) were similar to those obtained in control experiments with intact labyrinths. However, the mean discharge rate of the responsive units slightly decreased with respect to the value obtained in control experiments, the decrease being more prominent within the rvLVN. The average sensitivity (and to a lesser extent the gain) of responses of rvLVN neurons to the labyrinth input was almost twice that of the dcLVN units in preparations with the vestibular nerves intact; these regional differences disappeared after contralateral aVN, particularly due to a decrease in gain and sensitivity of responses in the rvLVN. The proportion of LVN neurons that were maximally excited by animal position increased from 74.0% in the control experiments to 82.8%. However, while in control experiments the proportion of units excited during side-down tilt was twice as high as that of the units excited by side-up tilt, the opposite occurred after contralateral aVN; this finding affected particularly the dcLVN. In addition the average phase lead of responses relative to the extreme animal displacements slightly decreased from +12.3 degrees in control experiments to +9.4 degrees. Among the LVN neurons recorded after contralateral aVN, 35 were antidromically activated by stimulating the spinal cord at T12 L1, while 43 units were not activated. The relation found in control experiments, i.e., that the faster the conduction velocity of vestibulospinal axon the lower was the unit discharge at rest, was lost after contralateral aVN, due to a decrease in resting discharge rate of the slow neurons. This finding, coupled with the observation that slow and fast units did not show any difference in their response gain to tilt, explains why the positive correlation between axonal conduction velocity and response sensitivity occurring in control experiments was lost after contralateral aVN.(ABSTRACT TRUNCATED AT 400 WORDS)

Abstract: The activity of 168 Deiters' neurons projecting to lumbosacral segments of the spinal cord has been recorded in precollicular decerebrate cats after ipsilateral acute (aVN) or chronic vestibular neurectomy (cVN), and their response characteristics to sinusoidal stimulation of contralateral labyrinth receptors at the standard parameters (roll tilt at 0.026 Hz, 10 degrees peak displacement) have been related to cell size inferred from the conduction velocity of the corresponding axons. These findings were compared with those elicited in decerebrate cats with both vestibular nerves intact. In all experimental conditions, the higher the coefficient of variation (CV) of the vestibulospinal neurons, reflecting a more irregular unit discharge, the lower was the mean discharge rate at rest. However, the proportion of regularly discharging units (with the lowest CV) decreased after aVN but increased after cVN. The relation found in control experiments, i.e., the faster the conduction velocity of vestibulospinal axon the lower was the unit discharge at rest, was lost after aVN due to a decrease in resting discharge rate of the slow neurons. The mean discharge rate of these units, however, recovered after cVN, so that the negative correlation between resting discharge rate and axonal conduction velocity was reestablished. After aVN, the decrease in resting discharge rate of the slow vestibulospinal neurons was not associated with significant changes in gain (impulses per second per degree) of the unit responses to standard parameters of tilt, so that the sensitivity of these units (percentage change of the mean discharge rate per degree) increased; on the other hand, the resting discharge rate of the fast neurons, which remained almost unchanged after aVN, was associated with a significant increase in gain, thus leading to an average increase in response sensitivity of these units.(ABSTRACT TRUNCATED AT 400 WORDS)

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Abstract: In the present investigation, we have analysed the visually induced modulations of muscular responses during falls at different rates of acceleration and performed in five different visual conditions: Normal vision (NV), Darkness (D), Stabilized vision (SV), with visual motion cues being enhanced (EV), or reduced (RV). This study was conducted on normal and hemilabyrinthectomized baboons. EMG activities were recorded in the alert monkey from three pairs of muscles (splenius capitis, soleus and tibialis anterior). For testing, the monkey was seated in a special chair unexpectedly dropped by 0.9 m. Five peaks of maximum acceleration were used (8.8, 6.6, 4.4, 3.3, 2.2 m/s2). Conditions EV, SV and RV were tested by way of projector, the input of which consisted of the integral of vertical acceleration and output, the output of which controlled film motion. In the normal baboon the visually induced modulation of EMG responses in the SV, EV, and RV conditions was larger for slow falls than for fast ones. This modulation was direction-specific, at least for slow falls, and depended on the relative speed of the visual scene. Between certain limits, the energy of the responses was roughly proportional to the relative speed of the visual scene. These modifications were most accentuated in the splenius and soleus muscles. Condition D only produced a slight reduction of the EMG response. All these findings eliminate the possibility that the observed effects represents a startle response. Thus, we can conclude that there is a fact directional role of vision in postural control in the normal falling baboon. In the hemilabyrinthectomized animal, greater modulations were recorded only when the visual manipulations were performed during the first two postoperative weeks. This confirms the above results on the normal baboon and previous data on the role of vision in the recovery process.

Abstract: In previous studies a contribution of vision to vestibular-dependent muscle responses during free-fall was found in the intact monkey, and the role of remaining labyrinthine afferents in compensation of these postural reactions was studied in vestibular neurectomized monkeys. In the present investigation we have compared the role of visual motion cues in the recovery of muscle responses to fall in unilateral (U.N.) and bilateral vestibular neurectomized (B.N.) baboons. During free-fall, electromyographic (EMG) responses were recorded from splenius capitis, soleus and tibialis anterior muscles. EMG activities were recorded in two randomly presented conditions: with normal motion of the visual world (NV) and with the visual world stabilized with respect to the baboon's head (SV) until 6 weeks after surgery. In B.N. baboons, results showed that condition SV was accompanied by a very strong motor depression during the entire test period. A greater decrease was observed in the splenius and soleus muscles. In U.N. baboons, significantly depressed EMG responses were recorded in the SV condition during the first two stages of compensation only (0--2 weeks), in all tested muscles except the tibialis anterior muscle. On the other hand, these motor depressions appeared to depend upon the level of neuronal resting activity in the vestibular nuclei. It is inferred that the partial recovery of muscle responses to fall observed in B.N. baboons in the NV condition is mainly due to visual information concerning motion, which replaces to the labyrinthine afferents. In U.N. baboons, the visual motion cues would fulfil only a transitory substitution function by supplying the decrease of neuronal activity in the vestibular nuclei. Later on, full compensation would be carried out by means of the remaining labyrinth.

Abstract: The role of sensorimotor activity in compensating deficits following unilateral vestibular neurectomy was studied in four adult cats using behavioral tests. Disturbances in posture and equilibrium were quantified and their subsequent compensation was described in both sensorimotor restrained and unrestrained cats. Sensorimotor restriction (S.M.R.) lasted 7 days and was performed in different postoperative periods. In the unrestrained animal, postural asymmetry compensation followed a 3-phase time course leading to preoperative criteria after about 40 days. Recovery of equilibrium developed by steps and was achieved after about 50 postoperative days. A first week applied S.M.R. was most effective in stopping postural symmetry recovery, while a later S.M.R. had no effect on the recovery time course and did not produce decompensation. On the contrary, S.M.R. (1st week or 3rd week) prevented and delayed equilibrium recovery, the earlier S.M.R. producing maximal effects. These observations suggest a CNS "sensitive period" to vicariant inputs.

Abstract: The electromyographic (EMG) responses from soleus and tibialis anterior muscles and the monosynaptic H- and T-reflex responses from soleus muscles were recorded bilaterally from conscious baboon while unexpectedly dropping it with unrestricted vision. These responses were recorded either after unilateral vestibular neurectomy (U.N. Baboons) or after bilateral neurectomy performed in one stage (B.N. 1 baboons) and in two stages (B.N. 2 baboons). A positive correlation was found between modifications and development of EMG responses and reflex data. In the U.N. baboons, some differences were observed when comparing data from the H- and T-reflex methods, suggesting that recovery of normal responses to fall is achieved both by means of direct influences on alpha-motoneurons and via the gamma-loop. In the U.N. baboons postural reactions to fall developed in three distinct periods. The first or critical stage showed asymmetrical EMG and reflex responses with increased responses from contralateral soleus muscle and decreased responses from ipsilateral soleus. Opposite effects were recorded from tibialis anterior flexor muscles. The second or acute stage which began around 4 to 7 days after surgery exhibited symmetrical, but very reduced, responses when compared to the control in soleus muscles, and symmetrical, but increased, responses from tibialis anterior muscles. This stage lasted until about the end of the second postoperative week and was followed by the third or compensatory stage during which EMG as well as reflex responses developed towards the control pattern in all tested muscles. Almost normal responses were recorded on both sides 3 weeks after surgery. Only a partial recovery was found in the B.N. 1 baboons, indicating that the contralateral remaining labyrinthine afferences constitute a necessary condition for the full compensation of postural reactions to fall in the case of unilateral vestibular neurectomy. The Bechterew's compensation was obtained in the B.N. 2 baboons. These results are discussed in relation with the general organization of the vestibulospinal pathways and with those concerning development of the postoperative activity at the vestibular nuclei level. A model of vestibular compensation achieved by means of a multisensory substitution process is suggested.

Abstract: The free fall has been used in our laboratory as a way to test vestibular function in baboons in order to quantify vestibular compensation in the hemilabyrinthectomized animal. This study presents only those results that concern the contribution of the vestibular system to muscle responses due to sudden fall. EMG activity was recorded from the fully conscious animal using chronic electrodes implanted in various muscles. Spinal monosynaptic reflexes (Hoffmann's and tendon reflexes) were studied in the soleus muscle. Baboons were seated in a special chair suspended from an electromagnet and unexpectedly dropped 90 cm. Experiments were performed in normal, unilateral and bilateral vestibular neurectomized baboons. 1. In normal baboons, results showed a first short-latency response in all tested muscles, followed by a second peak of EMG activity in these muscles. Comparison with data from bilateral vestibular neurectomized baboons demonstrates that normal vestibular function is essential for the appearance of the first peak; the second peak rapidly disappears in our experimental situation where the animal's fall is mechanically braked and interrupted, so the animal does not have to make the postural adjustments necessary for landing, It is suggested that the first peak is concerned with the automatic and reflex control of landing, the second with the voluntary breaking of landing. 2. The modulation of monosynaptic spinal reflexes is closely related to the EMG response in soleus muscle. Facilitation of the H-reflex begins just prior to the onset of the EMG activity and continues as long as the baboon is falling. The T-reflex modulation presents a similar time course except in its early phase where it is depressed. Decrease in T and increase in H-reflexes suggest that the EMG response is most likely due to direct activation of alpha-motoneurons and not by means of the gamma-loop. 3. In unilateral vestibular neurectomized baboons, EMG and reflexological data show the classical asymmetry characterized by a strong decrease of the responses on the side of the lesion, and by a pronounced increase on the contralateral side. It is concluded that this represents the imbalance between the resting discharge of the vestibular neurons, and discloses the influence of labyrinthine afferences at the spinal level. We suggest consequently the use of EMG responses and modulation of spinal reflexes to fall in order to quantify vestibular compensation.