RESUMEN
The dorsal root ganglia (DRG), housing primary sensory neurons, transmit somatosensory and visceral afferent inputs to the dorsal horn of the spinal cord. They play a pivotal role in both physiological and pathological states, including neuropathic and visceral pain. In vivo calcium imaging of DRG enables real-time observation of calcium transients in single units or neuron ensembles. Accumulating evidence indicates that DRG neuronal activities induced by somatic stimulation significantly affect autonomic and visceral functions. While lumbar DRG calcium imaging has been extensively studied, thoracic segment DRG calcium imaging has been less explored due to surgical exposure and stereotaxic fixation challenges. Here, we utilized in vivo calcium imaging at the thoracic1 dorsal root ganglion (T1-DRG) to investigate changes in neuronal activity resulting from somatic stimulations of the forelimb. This approach is crucial for understanding the somato-cardiac reflex triggered by peripheral nerve stimulations (PENS), such as acupuncture. Notably, synchronization of cardiac function was observed and measured by electrocardiogram (ECG), with T-DRG neuronal activities, potentially establishing a novel paradigm for somato-visceral reflex in the thoracic segments.
Asunto(s)
Calcio , Electrocardiografía , Ganglios Espinales , Animales , Ganglios Espinales/fisiología , Calcio/metabolismo , Calcio/análisis , Electrocardiografía/métodos , Ratones , Nervios Periféricos/fisiología , Miembro Anterior/inervación , Miembro Anterior/fisiologíaRESUMEN
BACKGROUND: Reaching and grasping (R&G) in rats is commonly used as an outcome measure to investigate the effectiveness of rehabilitation or treatment strategies to recover forelimb function post spinal cord injury. Kinematic analysis has been limited to the wrist and digit movements. Kinematic profiles of the more proximal body segments that play an equally crucial role in successfully executing the task remain unexplored. Additionally, understanding of different forelimb muscle activity, their interactions, and their correlation with the kinematics of R&G movement is scarce. NEW METHOD: In this work, novel methodologies to comprehensively assess and quantify the 3D kinematics of the proximal and distal forelimb joints along with associated muscle activity during R&G movements in adult rats are developed and discussed. RESULTS: Our data show that different phases of R&G identified using the novel kinematic and EMG-based approach correlate with the well-established descriptors of R&G stages derived from the Whishaw scoring system. Additionally, the developed methodology allows describing the temporal activity of individual muscles and associated mechanical and physiological properties during different phases of the motor task. COMPARISON WITH EXISTING METHOD(S): R&G phases and their sub-components are identified and quantified using the developed kinematic and EMG-based approach. Importantly, the identified R&G phases closely match the well-established qualitative descriptors of the R&G task proposed by Whishaw and colleagues. CONCLUSIONS: The present work provides an in-depth objective analysis of kinematics and EMG activity of R&G behavior, paving the way to a standardized approach to assessing this critical rodent motor function in future studies.
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Electromiografía , Miembro Anterior , Fuerza de la Mano , Músculo Esquelético , Animales , Fenómenos Biomecánicos/fisiología , Miembro Anterior/fisiología , Electromiografía/métodos , Músculo Esquelético/fisiología , Fuerza de la Mano/fisiología , Ratas , Traumatismos de la Médula Espinal/fisiopatología , Femenino , Destreza Motora/fisiología , Masculino , Ratas Sprague-Dawley , Conducta Animal/fisiología , Movimiento/fisiologíaRESUMEN
Coordinated movement requires the nervous system to continuously compensate for changes in mechanical load across different conditions. For voluntary movements like reaching, the motor cortex is a critical hub that generates commands to move the limbs and counteract loads. How does cortex contribute to load compensation when rhythmic movements are sequenced by a spinal pattern generator? Here, we address this question by manipulating the mass of the forelimb in unrestrained mice during locomotion. While load produces changes in motor output that are robust to inactivation of motor cortex, it also induces a profound shift in cortical dynamics. This shift is minimally affected by cerebellar perturbation and significantly larger than the load response in the spinal motoneuron population. This latent representation may enable motor cortex to generate appropriate commands when a voluntary movement must be integrated with an ongoing, spinally-generated rhythm.
Asunto(s)
Locomoción , Corteza Motora , Neuronas Motoras , Animales , Corteza Motora/fisiología , Ratones , Neuronas Motoras/fisiología , Locomoción/fisiología , Miembro Anterior/fisiología , Masculino , Ratones Endogámicos C57BL , Movimiento/fisiología , Médula Espinal/fisiología , Femenino , Cerebelo/fisiologíaRESUMEN
Take-off is a vital part of powered flight which likely constrains the size of birds, yet extinct pterosaurs are known to have reached far larger sizes. Three different hypothesised take-off motions (bipedal burst launching, bipedal countermotion launching, and quadrupedal launching) have been proposed as explanations for how pterosaurs became airborne and circumvented this proposed morphological limit. We have constructed a computational musculoskeletal model of a 5 m wingspan ornithocheiraean pterosaur, reconstructing thirty-four key muscles to estimate the muscle moment arms throughout the three hypothesised take-off motions. Range of motion constrained hypothetical kinematic sequences for bipedal and quadrupedal take-off motions were modelled after extant flying vertebrates. Across our simulations we did not find higher hindlimb moment arms for bipedal take-off motions or noticeably higher forelimb moment arms in the forelimb for quadrupedal take-off motions. Despite this, in all our models we found the muscles utilised in the quadrupedal take-off have the largest total launch applicable moment arms throughout the entire take-off sequences and for the take-off pose. This indicates the potential availability of higher leverage for a quadrupedal take-off than hypothesised bipedal motions in pterosaurs pending further examination of muscle forces.
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Dinosaurios , Vuelo Animal , Animales , Fenómenos Biomecánicos , Vuelo Animal/fisiología , Dinosaurios/fisiología , Dinosaurios/anatomía & histología , Músculo Esquelético/fisiología , Músculo Esquelético/anatomía & histología , Fósiles , Modelos Biológicos , Miembro Anterior/fisiología , Miembro Anterior/anatomía & histología , Simulación por ComputadorRESUMEN
Hoof care providers are pivotal for implementing biomechanical optimizations of the musculoskeletal system in the horse. Regular visits allow for the collection of longitudinal, quantitative information ("normal ranges"). Changes in movement symmetry, e.g., after shoeing, are indicative of alterations in weight-bearing and push-off force production. Ten Warmblood show jumping horses (7-13 years; 7 geldings, 3 mares) underwent forelimb re-shoeing with rolled rocker shoes, one limb at a time ("limb-by-limb"). Movement symmetry was measured with inertial sensors attached to the head, withers, and pelvis during straight-line trot and lunging. Normalized differences pre/post re-shoeing were compared to published test-retest repeatability values. Mixed-model analysis with random factors horse and limb within horse and fixed factors surface and exercise direction evaluated movement symmetry changes (p < 0.05, Bonferroni correction). Withers movement indicated increased forelimb push-off with the re-shod limb on the inside of the circle and reduced weight-bearing with the re-shod limb and the ipsilateral hind limb on hard ground compared to soft ground. Movement symmetry measurements indicate that a rolled rocker shoe allows for increased push-off on soft ground in trot in a circle. Similar studies should study different types of shoes for improved practically relevant knowledge about shoeing mechanics, working towards evidence-based preventative shoeing.
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Miembro Anterior , Zapatos , Animales , Caballos/fisiología , Miembro Anterior/fisiología , Fenómenos Biomecánicos/fisiología , Movimiento/fisiología , Soporte de Peso/fisiología , Marcha/fisiología , Femenino , Masculino , Miembro Posterior/fisiologíaRESUMEN
Hominoids exhibit a strikingly diverse set of locomotor adaptations-including knuckle-walking, brachiation, quadrumanuous suspension, and striding bipedalism-while also possessing morphologies associated with forelimb suspension. It has been suggested that changes in limb element integration facilitated the evolution of diverse locomotor modes by reducing covariation between serial homologs and allowing the evolution of a greater diversity of limb lengths. Here, I compare limb element integration in hominoids with that of other primate taxa, including two that have converged with them in forelimb morphology, Ateles and Pygathrix. Ateles is part of a clade that, such as hominoids, exhibits diverse locomotor adaptations, whereas Pygathrix is an anomaly in a much more homogeneous (in terms of locomotor adaptations) clade. I find that all atelines (and possibly all atelids), not just Ateles, share reduced limb element integration with hominoids. Pygathrix does not, however, instead resembling other members of its own family. Indriids also seem to have higher limb integration than apes, despite using their forelimbs and hindlimbs in divergent ways, although there is more uncertainty in this group due to poor sample size. These results suggest that reduced limb integration is characteristic of certain taxonomic groups with high locomotor diversity rather than taxa with specific, specialized locomotor adaptations. This is consistent with the hypothesis that reduced integration serves to open new areas of morphospace to those clades while suggesting that derived locomotion with divergent demands on limbs is not necessarily associated with reduced limb integration.
Asunto(s)
Locomoción , Animales , Evolución Biológica , Adaptación Fisiológica , Fósiles/anatomía & histología , Primates/fisiología , Primates/anatomía & histología , Extremidades/anatomía & histología , Extremidades/fisiología , Miembro Anterior/anatomía & histología , Miembro Anterior/fisiologíaRESUMEN
Holding still and aiming reaches to spatial targets may depend on distinct neural circuits. Using automated homecage training and a sensitive joystick, we trained freely moving mice to contact a joystick, hold their forelimb still, and then reach to rewarded target locations. Mice learned the task by initiating forelimb sequences with clearly resolved submillimeter-scale micromovements followed by millimeter-scale reaches to learned spatial targets. Hundreds of thousands of trajectories were decomposed into millions of kinematic submovements, while photoinhibition was used to test roles of motor cortical areas. Inactivation of both caudal and rostral forelimb areas preserved the ability to produce aimed reaches, but reduced reach speed. Inactivation specifically of contralateral caudal forelimb area (CFA) additionally impaired the ability to aim corrective submovements to remembered locations following target undershoots. Our findings show that motor cortical inactivations reduce the gain of forelimb movements but that inactivation specifically of contralateral CFA impairs corrective movements important for reaching a target location.NEW & NOTEWORTHY To test the role of different cortical areas in holding still and reaching to targets, this study combined home-cage training with optogenetic silencing as mice engaged in a learned center-out-reach task. Inactivation specifically of contralateral caudal forelimb area (CFA) impaired corrective movements necessary to reach spatial targets to earn reward.
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Miembro Anterior , Corteza Motora , Desempeño Psicomotor , Animales , Corteza Motora/fisiología , Ratones , Miembro Anterior/fisiología , Masculino , Desempeño Psicomotor/fisiología , Ratones Endogámicos C57BL , Optogenética , Fenómenos Biomecánicos , Femenino , Movimiento/fisiología , Actividad Motora/fisiologíaRESUMEN
Bats are the only mammals capable of powered flight and have correspondingly specialized body plans, particularly in their limb morphology. The origin of bat flight is still not fully understood due to an uninformative fossil record but, from the perspective of a functional transition, it is widely hypothesized that bats evolved from gliding ancestors. Here, we test predictions of the gliding-to-flying hypothesis of the origin of bat flight by using phylogenetic comparative methods to model the evolution of forelimb and hindlimb traits on a dataset spanning four extinct bats and 231 extant mammals with diverse locomotor modes. Our results reveal that gliders exhibit adaptive trait optima (1) toward relatively elongate forelimbs that are intermediate between those of bats and non-gliding arborealists, and (2) toward relatively narrower but not longer hindlimbs that are intermediate between those of non-gliders and bats. We propose an adaptive landscape based on limb length and width optimal trends derived from our modeling analyses. Our results support a hypothetical evolutionary pathway wherein glider-like postcranial morphology precedes a bat-like morphology adapted to powered-flight, setting a foundation for future developmental, biomechanical, and evolutionary research to test this idea.
Asunto(s)
Evolución Biológica , Quirópteros , Vuelo Animal , Miembro Anterior , Filogenia , Quirópteros/anatomía & histología , Quirópteros/fisiología , Animales , Vuelo Animal/fisiología , Miembro Anterior/anatomía & histología , Miembro Anterior/fisiología , Miembro Posterior/anatomía & histología , Miembro Posterior/fisiología , Fósiles , Fenómenos BiomecánicosRESUMEN
Equine physiotherapy commonly includes basic exercises such as walking backward (BW) and voluntary lifting of single limbs (SLL), but trunk movements during these have not been studied. In order to compare the trunk kinematics during BW and SLL with forward walking (FW), nine horses were measured in FW, BW and during SLL triggered by tactile cue. Kinematics were obtained from skin markers captured by ten high-speed video cameras. Trunk angles were calculated in sagittal and horizontal planes from withers, dorsal to spinous processes of the 16th thoracic vertebra (T16), 2nd and 4th sacral vertebrae (S2, S4), WT16S2 and T16S2S4 respectively. From the hooves, maximum hoof height during swing phase and horizontal distance between hoof and median body plane during swing and stance phases were determined. Dorsoventral range of motion (ROM) and maximum flexion of WT16S2 was significantly larger in BW than in FW, while laterolateral ROM was significantly smaller during hindlimb swing phase in BW and SLL than in FW. In contrast, dorsoventral ROM of T16S2S4 was significantly smaller during stance and swing phases of hindlimbs in BW compared to FW, and throughout the movement. During forelimb swing phase, T16S2S4 ROM was significantly larger in BW than SLL. Hindhoof height in SLL was significantly higher than in FW. Distance between median body plane and hooves was significantly larger in BW than in FW, and significantly larger in BW than in SLL for hindlimb swing phase. In BW, increased lumbosacral stabilisation and the larger area of support created by fore- and hindlimbs may represent a strategy to enhance body stabilisation, as BW entails some insecurity.
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Miembro Anterior , Miembro Posterior , Caminata , Animales , Caballos/fisiología , Fenómenos Biomecánicos , Miembro Posterior/fisiología , Miembro Anterior/fisiología , Caminata/fisiología , Masculino , Femenino , Marcha/fisiología , Torso/fisiología , Rango del Movimiento ArticularRESUMEN
Whether the forelimb-digging apparatus of tooth-digging subterranean mammals has similar levels of specialization as compared to scratch-diggers is still unknown. We assessed the scapular morphology and forelimb musculature of all four solitary African mole rats (Bathyergidae): two scratch-diggers, Bathyergus suillus and Bathyergus janetta, and two chisel-tooth diggers, Heliophobius argenteocinereus and Georychus capensis. Remarkable differences were detected: Bathyergus have more robust neck, shoulder, and forearm muscles as compared to the other genera. Some muscles in Bathyergus were also fused and often showing wider attachment areas to bones, which correlate well with its more robust and larger scapula, and its wider and medially oriented olecranon. This suggests that shoulder, elbow, and wrist work in synergy in Bathyergus for generating greater out-forces and that the scapula and proximal ulna play fundamental roles as pivots to maximize and accommodate specialized muscles for better (i) glenohumeral and scapular stabilization, (ii) powerful shoulder flexion, (iii) extension of the elbow and (iv) flexion of the manus and digits. Moreover, although all bathyergids showed a similar set of muscles, Heliophobius lacked the m. tensor fasciae antebrachii (aiding with elbow extension and humeral retraction), and Heliophobius and Georychus lacked the m. articularis humeri (aiding with humeral adduction), indicating deeper morphogenetic differences among digging groups and suggesting a relatively less specialized scratch-digging ability. Nevertheless, Heliophobius and Bathyergus shared some similar adaptations allowing scratch-digging. Our results provide new information about the morphological divergence within this family associated with the specialization to distinct functions and digging behaviors, thus contributing to understand the mosaic of adaptations emerging in phylogenetically and ecologically closer subterranean taxa. This and previous anatomical studies on the Bathyergidae will provide researchers with a substantial basis on the form and function of the musculoskeletal system for future kinematic investigations of digging behavior, as well as to define potential indicators of scratch-digging ability.
Asunto(s)
Miembro Anterior , Animales , Miembro Anterior/anatomía & histología , Miembro Anterior/fisiología , Músculo Esquelético/anatomía & histología , Músculo Esquelético/fisiología , Ratas Topo/anatomía & histología , Ratas Topo/fisiología , Sistema Musculoesquelético/anatomía & histología , Escápula/anatomía & histología , Escápula/fisiologíaRESUMEN
During quadrupedal locomotion, interactions between spinal and supraspinal circuits and somatosensory feedback coordinate forelimb and hindlimb movements. How this is achieved is not clear. To determine whether forelimb movements modulate hindlimb cutaneous reflexes involved in responding to an external perturbation, we stimulated the superficial peroneal nerve in six intact cats during quadrupedal locomotion and during hindlimb-only locomotion (with forelimbs standing on stationary platform) and in two cats with a low spinal transection (T12-T13) during hindlimb-only locomotion. We compared cutaneous reflexes evoked in six ipsilateral and four contralateral hindlimb muscles. Results showed similar occurrence and phase-dependent modulation of short-latency inhibitory and excitatory responses during quadrupedal and hindlimb-only locomotion in intact cats. However, the depth of modulation was reduced in the ipsilateral semitendinosus during hindlimb-only locomotion. Additionally, longer-latency responses occurred less frequently in extensor muscles bilaterally during hindlimb-only locomotion, whereas short-latency inhibitory and longer-latency excitatory responses occurred more frequently in the ipsilateral and contralateral sartorius anterior, respectively. After spinal transection, short-latency inhibitory and excitatory responses were similar to both intact conditions, whereas mid- or longer-latency excitatory responses were reduced or abolished. Our results in intact cats and the comparison with spinal-transected cats suggest that the absence of forelimb movements suppresses inputs from supraspinal structures and/or cervical cord that normally contribute to longer-latency reflex responses in hindlimb extensor muscles.NEW & NOTEWORTHY During quadrupedal locomotion, the coordination of forelimb and hindlimb movements involves central circuits and somatosensory feedback. To demonstrate how forelimb movement affects hindlimb cutaneous reflexes during locomotion, we stimulated the superficial peroneal nerve in intact cats during quadrupedal and hindlimb-only locomotion as well as in spinal-transected cats during hindlimb-only locomotion. We show that forelimb movement influences the modulation of hindlimb cutaneous reflexes, particularly the occurrence of long-latency reflex responses.
Asunto(s)
Miembro Anterior , Miembro Posterior , Locomoción , Músculo Esquelético , Reflejo , Traumatismos de la Médula Espinal , Animales , Gatos , Miembro Posterior/fisiología , Miembro Anterior/fisiología , Reflejo/fisiología , Locomoción/fisiología , Músculo Esquelético/fisiología , Traumatismos de la Médula Espinal/fisiopatología , Movimiento/fisiología , Femenino , Masculino , Piel/inervaciónRESUMEN
The pedunculopontine tegmental nucleus of the brainstem (PPTg) has extensive interconnections and neuronal-behavioural correlates. It is implicated in movement control and sensorimotor integration. We investigated whether single neuron activity in freely moving rats is correlated with components of skilled forelimb movement, and whether individual neurons respond to both motor and sensory events. We found that individual PPTg neurons showed changes in firing rate at different times during the reach. This type of temporally specific modulation is like activity seen elsewhere in voluntary movement control circuits, such as the motor cortex, and suggests that PPTg neural activity is related to different specific events occurring during the reach. In particular, many neuronal modulations were time-locked to the end of the extension phase of the reach, when fine distal movements related to food grasping occur, indicating strong engagement of PPTg in this phase of skilled individual forelimb movements. In addition, some neurons showed brief periods of apparent oscillatory firing in the theta range at specific phases of the reach-to-grasp movement. When movement-related neurons were tested with tone stimuli, many also responded to this auditory input, allowing for sensorimotor integration at the cellular level. Together, these data extend the concept of the PPTg as an integrative structure in generation of complex movements, by showing that this function extends to the highly coordinated control of the forelimb during skilled reach to grasp movement, and that sensory and motor-related information converges on single neurons, allowing for direct integration at the cellular level.
Asunto(s)
Neuronas , Núcleo Tegmental Pedunculopontino , Ritmo Teta , Animales , Núcleo Tegmental Pedunculopontino/fisiología , Neuronas/fisiología , Ratas , Masculino , Ritmo Teta/fisiología , Movimiento/fisiología , Miembro Anterior/fisiología , Ratas Long-Evans , Potenciales de Acción/fisiología , Estimulación Acústica/métodosRESUMEN
Rhythmic movement is the fundamental motion dynamics characterized by repetitive patterns. Precisely defining onsets in rhythmic movement is essential for a comprehensive analysis of motor functions. Our study introduces an automated method for detecting rat's forelimb foot-strike onsets using deep learning tools. This method demonstrates high accuracy of onset detection by combining two techniques using joint coordinates and behavioral confidence scale. The analysis extends to neural oscillatory responses in the rat's somatosensory cortex, validating the effectiveness of our combined approach. Our technique streamlines experimentation, demanding only a camera and GPU-accelerated computer. This approach is applicable across various contexts and promotes our understanding of brain functions during rhythmic movements.
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Pie , Miembro Anterior , Movimiento , Corteza Somatosensorial , Animales , Miembro Anterior/fisiología , Movimiento/fisiología , Ratas , Corteza Somatosensorial/fisiología , Pie/fisiología , Masculino , Periodicidad , Aprendizaje ProfundoRESUMEN
Although active touch in rodents arises from the forepaws as well as whiskers, most research on active touch only focuses on whiskers. This results in a paucity of tasks designed to assess the process of active touch with a forepaw. We develop a new experimental task, the Reach-to-Grasp and Tactile Discrimination task (RGTD task), to examine active touch with a forepaw in rodents, particularly changes in processes of active touch during motor skill learning. In the RGTD task, animals are required to (1) extend their forelimb to an object, (2) grasp the object, and (3) manipulate the grasped object with the forelimb. The animals must determine the direction of the manipulation based on active touch sensations arising during the period of the grasping. In experiment 1 of the present study, we showed that rats can learn the RGTD task. In experiment 2, we confirmed that the rats are capable of reversal learning of the RGTD task. The RGTD task shared most of the reaching movements involved with conventional forelimb reaching tasks. From the standpoint of a discrimination task, the RGTD task enables rigorous experimental control, for example by removing bias in the stimulus-response correspondence, and makes it possible to utilize diverse experimental procedures that have been difficult in prior tasks.
Asunto(s)
Aprendizaje Discriminativo , Miembro Anterior , Tacto , Animales , Ratas , Masculino , Miembro Anterior/fisiología , Tacto/fisiología , Aprendizaje Discriminativo/fisiología , Fuerza de la Mano/fisiología , Percepción del Tacto/fisiología , Desempeño Psicomotor/fisiología , Discriminación en Psicología/fisiología , Destreza Motora/fisiología , Ratas Long-Evans , Aprendizaje Inverso/fisiologíaRESUMEN
OBJECTIVE: To perform non-invasive Electroarthrography (EAG) on live horses and establish relationships between EAG and direct measurements of cartilage streaming potentials in weight bearing areas of the equine metacarpophalangeal joint. DESIGN: EAG was performed bilaterally on the metacarpophalangeal joints of live horses (n = 3). Separate experiments used metacarpophalangeal joint explants (n = 11) to measure EAG obtained during simulated loading followed by direct measurements of cartilage streaming potentials on joint surfaces using the Arthro-BST probe. Joints were assigned to relatively normal (n = 5) and mildly degraded (n = 6) groups based on histological scoring of Safranin-O/Fast Green stained sections. RESULTS: EAG, involving application of electrodes to skin surrounding the joint and repeated weight shifting, was well-tolerated in live horses. One pair of distal forelimbs were available for analogous ex vivo EAG testing and measurements were strongly correlated to in vivo EAG measurements obtained on the same joints (r = 0.804, p = 0.016, n = 8). Both indirect (EAG) and direct (Arthro-BST) measurements of cartilage streaming potentials distinguished between normal and mildly degraded cartilage with statistically significant differences at 5 of 6 and 4 of 6 electrodes during simulated standing and walking, respectively. Strong and moderate correlations for weight bearing regions on the dorsal phalanx and central metacarpus were detected during both standing and walking. At the metacarpus/sesamoid interface a moderate correlation occurred during walking. CONCLUSION: Non-invasive EAG was used successfully in a clinical scenario and correlated to direct measurements of streaming potentials in weight bearing cartilage. These data support the potential of EAG to contribute to the diagnosis and treatment of degenerative joint diseases.
Asunto(s)
Cartílago Articular , Articulación Metacarpofalángica , Soporte de Peso , Animales , Caballos , Cartílago Articular/diagnóstico por imagen , Cartílago Articular/fisiología , Articulación Metacarpofalángica/diagnóstico por imagen , Articulación Metacarpofalángica/fisiología , Articulación Metacarpofalángica/fisiopatología , Soporte de Peso/fisiología , Artrografía/métodos , Miembro Anterior/fisiologíaRESUMEN
When the foot dorsum contacts an obstacle during locomotion, cutaneous afferents signal central circuits to coordinate muscle activity in the four limbs. Spinal cord injury disrupts these interactions, impairing balance and interlimb coordination. We evoked cutaneous reflexes by electrically stimulating left and right superficial peroneal nerves before and after two thoracic lateral hemisections placed on opposite sides of the cord at 9- to 13-week interval in seven adult cats (4 males and 3 females). We recorded reflex responses in ten hindlimb and five forelimb muscles bilaterally. After the first (right T5-T6) and second (left T10-T11) hemisections, coordination of the fore- and hindlimbs was altered and/or became less consistent. After the second hemisection, cats required balance assistance to perform quadrupedal locomotion. Short-latency reflex responses in homonymous and crossed hindlimb muscles largely remained unaffected after staggered hemisections. However, mid- and long-latency homonymous and crossed responses in both hindlimbs occurred less frequently after staggered hemisections. In forelimb muscles, homolateral and diagonal mid- and long-latency response occurrence significantly decreased after the first and second hemisections. In all four limbs, however, when present, short-, mid- and long-latency responses maintained their phase-dependent modulation. We also observed reduced durations of short-latency inhibitory homonymous responses in left hindlimb extensors early after the first hemisection and delayed short-latency responses in the right ipsilesional hindlimb after the first hemisection. Therefore, changes in cutaneous reflex responses correlated with impaired balance/stability and interlimb coordination during locomotion after spinal cord injury. Restoring reflex transmission could be used as a biomarker to facilitate locomotor recovery. KEY POINTS: Cutaneous afferent inputs coordinate muscle activity in the four limbs during locomotion when the foot dorsum contacts an obstacle. Thoracic spinal cord injury disrupts communication between spinal locomotor centres located at cervical and lumbar levels, impairing balance and limb coordination. We investigated cutaneous reflexes during quadrupedal locomotion by electrically stimulating the superficial peroneal nerve bilaterally, before and after staggered lateral thoracic hemisections of the spinal cord in cats. We showed a loss/reduction of mid- and long-latency responses in all four limbs after staggered hemisections, which correlated with altered coordination of the fore- and hindlimbs and impaired balance. Targeting cutaneous reflex pathways projecting to the four limbs could help develop therapeutic approaches aimed at restoring transmission in ascending and descending spinal pathways.
Asunto(s)
Miembro Posterior , Locomoción , Músculo Esquelético , Reflejo , Traumatismos de la Médula Espinal , Animales , Gatos , Miembro Posterior/inervación , Miembro Posterior/fisiología , Miembro Posterior/fisiopatología , Masculino , Femenino , Traumatismos de la Médula Espinal/fisiopatología , Reflejo/fisiología , Locomoción/fisiología , Músculo Esquelético/inervación , Músculo Esquelético/fisiología , Músculo Esquelético/fisiopatología , Piel/inervación , Vértebras Torácicas , Miembro Anterior/fisiopatología , Miembro Anterior/fisiología , Estimulación EléctricaRESUMEN
Layer 5 neurons of the neocortex receive their principal inputs from layer 2/3 neurons. We seek to identify the nature and extent of the plasticity of these projections with motor learning. Using optogenetic and viral intersectional tools to selectively stimulate distinct neuronal subsets in rat primary motor cortex, we simultaneously record from pairs of corticospinal neurons associated with distinct features of motor output control: distal forelimb vs. proximal forelimb. Activation of Channelrhodopsin2-expressing layer 2/3 afferents onto layer 5 in untrained animals produces greater monosynaptic excitation of neurons controlling the proximal forelimb. Following skilled grasp training, layer 2/3 inputs onto corticospinal neurons controlling the distal forelimb associated with skilled grasping become significantly stronger. Moreover, peak excitatory response amplitude nearly doubles while latency shortens, and excitatory-to-inhibitory latencies become significantly prolonged. These findings demonstrate distinct, highly segregated, and cell-specific plasticity of layer 2/3 projections during skilled grasp motor learning.
Asunto(s)
Miembro Anterior , Corteza Motora , Plasticidad Neuronal , Animales , Miembro Anterior/fisiología , Plasticidad Neuronal/fisiología , Corteza Motora/fisiología , Corteza Motora/citología , Ratas , Aprendizaje/fisiología , Fuerza de la Mano/fisiología , Neuronas/fisiología , Masculino , Tractos Piramidales/fisiología , Destreza Motora/fisiología , Femenino , Optogenética , Ratas Long-EvansRESUMEN
Heavy animals incur large forces on their limb bones, due to the transmission of body weight and ground reaction forces, and the contractions of the various muscles of the limbs. This is particularly true for rhinoceroses, the heaviest extant animals capable of galloping. Several studies have examined their musculoskeletal system and the forces their bones incur, but no detailed quantification has ever been attempted. Such quantification could help understand better the link between form and function in giant land animals. Here we constructed three-dimensional musculoskeletal models of the forelimb and hindlimb of Ceratotherium simum, the heaviest extant rhino species, and used static optimisation (inverse) simulations to estimate the forces applied on the bones when standing at rest, including magnitudes and directions. Overall, unsurprisingly, the most active muscles were antigravity muscles, which generate moments opposing body weight (thereby incurring the ground reaction force), and thus keep the joints extended, avoiding joint collapse via flexion. Some muscles have an antigravity action around several joints, and thus were found to be highly active, likely specialised in body weight support (ulnaris lateralis; digital flexors). The humerus was subjected to the greatest amount of forces in terms of total magnitude; forces on the humerus furthermore came from a great variety of directions. The radius was mainly subject to high-magnitude compressive joint reaction forces, but to little muscular tension, whereas the opposite pattern was observed for the ulna. The femur had a pattern similar to that of the humerus, and the tibia's pattern was intermediate, being subject to great compression in its caudal side but to great tension in its cranial side (i.e. bending). The fibula was subject to by far the lowest force magnitude. Overall, the forces estimated were consistent with the documented morphofunctional adaptations of C. simum's long bones, which have larger insertion areas for several muscles and a greater robusticity overall than those of lighter rhinos, likely reflecting the intense forces we estimated here. Our estimates of muscle and bone (joint) loading regimes for this giant tetrapod improve the understanding of the links between form and function in supportive tissues and could be extended to other aspects of bone morphology, such as microanatomy.
Asunto(s)
Músculo Esquelético , Perisodáctilos , Animales , Perisodáctilos/fisiología , Perisodáctilos/anatomía & histología , Fenómenos Biomecánicos/fisiología , Músculo Esquelético/fisiología , Músculo Esquelético/anatomía & histología , Miembro Anterior/fisiología , Miembro Anterior/anatomía & histología , Miembro Posterior/fisiología , Miembro Posterior/anatomía & histología , Simulación por ComputadorRESUMEN
Recent evidence suggests that primary sensory cortical regions play a role in the integration of information from multiple sensory modalities. How primary cortical neurons integrate different sources of sensory information is unclear, partly because non-primary sensory input to a cortical sensory region is often weak or modulatory. To address this question, we take advantage of the robust representation of thermal (cooling) and tactile stimuli in mouse forelimb primary somatosensory cortex (fS1). Using a thermotactile detection task, we show that the perception of threshold-level cool or tactile information is enhanced when they are presented simultaneously, compared with presentation alone. To investigate the cortical cellular correlates of thermotactile integration, we performed in vivo extracellular recordings from fS1 in awake resting and anesthetized mice during unimodal and bimodal stimulation of the forepaw. Unimodal stimulation evoked thermal- or tactile- specific excitatory and inhibitory responses of fS1 neurons. The most prominent features of combined thermotactile stimulation are the recruitment of unimodally silent fS1 neurons, non-linear integration features, and response dynamics that favor longer response durations with additional spikes. Together, we identify quantitative and qualitative changes in cortical encoding that may underlie the improvement in perception of thermotactile surfaces during haptic exploration.
Asunto(s)
Corteza Somatosensorial , Animales , Ratones , Corteza Somatosensorial/fisiología , Tacto/fisiología , Neuronas/fisiología , Ratones Endogámicos C57BL , Miembro Anterior/fisiología , Percepción del Tacto/fisiología , Masculino , Estimulación FísicaRESUMEN
The brain can generate actions, such as reaching to a target, using different movement strategies. We investigate how such strategies are learned in a task where perched head-fixed mice learn to reach to an invisible target area from a set start position using a joystick. This can be achieved by learning to move in a specific direction or to a specific endpoint location. As mice learn to reach the target, they refine their variable joystick trajectories into controlled reaches, which depend on the sensorimotor cortex. We show that individual mice learned strategies biased to either direction- or endpoint-based movements. This endpoint/direction bias correlates with spatial directional variability with which the workspace was explored during training. Model-free reinforcement learning agents can generate both strategies with similar correlation between variability during training and learning bias. These results provide evidence that reinforcement of individual exploratory behavior during training biases the reaching strategies that mice learn.