In this essay we describe some of the brain processes that are engaged when artists produce drawings and paintings. Two brain structures have been recently identified as key locations where the abstract neural signals representing the intention to move are transformed into patterns of neural activity suitable for activating the muscles. The first of these locations is the putamen, a section of the basal ganglia, the second is in the spinal cord. It is interesting that different types of modules and combinatorial activities characterize both structures. Finally, we discuss the ability of experts to generalize their learning across radically different contexts.
How is an evanescent wish to move translated into a concrete action? This simple question and puzzling miracle remains a focal point of motor systems neuroscience. Where does the difficulty lie? A great deal has been known about biomechanics for quite some time. More recently, there have been significant advances in our understanding of how the spinal system is organized into modules corresponding to spinal synergies, which are fixed patterns of multimuscle recruitment. But much less is known about how the supraspinal system recruits these synergies in the correct spatiotemporal pattern to effectively control movement. We argue that what makes the problem of supraspinal control so difficult is that it emerges as a result of multiple convergent and redundant sensorimotor loops. Because these loops are convergent, multiple modes of information are mixed before being sent to the spinal system; because they are redundant, information is overlapping such that a mechanism must exist to eliminate the redundancy before the signal is sent to the spinal system. Given these complex interactions, simple correlation analyses between movement variables and neural activity are likely to render a confusing and inconsistent picture. Here, we suggest that the perspective of sensorimotor loops might help in achieving a better systems-level understanding. Furthermore, state-of-the-art techniques in neurotechnology, such as optogenetics, appear to be well suited for investigating the problem of motor control at the level of loops.
The development of surface ElectroMyoGraphy (sEMG) acquisitionRossi, Fabio system having an optimal trade-off betweenMotto Ros, Paolo accuracy, resolution, low dimension and power consumptionSapienza, Stefano is a hot topic today. The event-driven Average Threshold Crossing (ATC) technique applied to the sEMG signal allows theBonato, Paolo reduction of both complexity and power consumption of the acquisition board. The paper presents an sEMG acquisitionBizzi, Emilio system, based on this approach, and shows the advantages of the ATC in this field. A framework for developing bio-signal ATC-processing applications is provided, enabling the comparisonDemarchi, Danilo with a standard sEMG sampling approach. Both system performance and power consumption analyses are carried out to obtain promising results in terms of real-time behavior and energy saving. As a sample application, the system is employed in the control of Functional Electrical Stimulation (FES) in way to verify the behavior of the ATC approach in such application.
In this paper some of the theoretical and experimental approaches related to multijoint arm trajectory formation are discussed. The problems I have considered are (a) coordinate transformation underlying trajectory planning , (b) the approaches elaborated by computer scientists to execute movements in artificial multijoint systems, and (c) a biological perspective on the problem of computing torques with muscles.
If speech is one possible sensorimotor instantiation of a more universal mental propensity, then why throughout human history has the spoken word been preferred to, say, the signed word?
The central pattern generator (CPG) architecture for rhythm generation remains partly elusive. We compare cat and frog locomotion results, where the component unrelated to pattern formation appears as a temporal grid, and traveling wave respectively. Frog spinal cord microstimulation with N-methyl-D-Aspartate (NMDA), a CPG activator, produced a limited set of force directions, sometimes tonic, but more often alternating between directions similar to the tonic forces. The tonic forces were topographically organized, and sites evoking rhythms with different force subsets were located close to the constituent tonic force regions. Thus CPGs consist of topographically organized modules. Modularity was also identified as a limited set of muscle synergies whose combinations reconstructed the EMGs. The cat CPG was investigated using proprioceptive inputs during fictive locomotion. Critical points identified both as abrupt transitions in the effect of phasic perturbations, and burst shape transitions, had biomechanical correlates in intact locomotion. During tonic proprioceptive perturbations, discrete shifts between these critical points explained the burst durations changes, and amplitude changes occurred at one of these points. Besides confirming CPG modularity, these results suggest a fixed temporal grid of anchoring points, to shift modules onsets and offsets. Frog locomotion, reconstructed with the NMDA synergies, showed a partially overlapping synergy activation sequence. Using the early synergy output evoked by NMDA at different spinal sites, revealed a rostrocaudal topographic organization, where each synergy is preferentially evoked from a few, albeit overlapping, cord regions. Comparing the locomotor synergy sequence with this topography suggests that a rostrocaudal traveling wave would activate the synergies in the proper sequence for locomotion. This output was reproduced in a two-layer model using this topography and a traveling wave. Together our results suggest two CPG components: modules, i.e., synergies; and temporal patterning, seen as a temporal grid in the cat, and a traveling wave in the frog. Animal and limb navigation have similarities. Research relating grid cells to the theta rhythm and on segmentation during navigation may relate to our temporal grid and traveling wave results. Winfree’s mathematical work, combining critical phases and a traveling wave, also appears important. We conclude suggesting tracing, and imaging experiments to investigate our CPG model.
Background Persons with Multiple Sclerosis frequently have gait deficits that lead to diminished activities of daily living. Identification of motoneuron activity patterns may elucidate new insight into impaired locomotor coordination and underlying neural systems. The aim of the present study was to investigate muscle synergies, identified by motor modules and their activation profiles, in persons with Multiple Sclerosis (PwMS) during walking compared to those of healthy subjects (HS), as well as, exploring relationship of muscle synergies with walking ability of PwMS. Methods Seventeen PwMS walked at their natural speed while 12 HS walked at slower than their natural speeds in order to provide normative gait values at matched speeds (spatio-temporal, kinematic and kinetic parameters and electromyography signals). Non-negative matrix factorization was used to identify muscle synergies from eight muscles. Pearson's correlation coefficient was used to evaluate the similarity of motor modules between PwMS and HS. To assess differences in module activations, each module's activation timing was integrated over 100% of gait cycle and the activation percentage was computed in six phases. Results Fifty-nine% of PwMS and 58% of HS had 4 modules while the remaining of both populations had 3 modules. Module 2 (related to soleus, medial and lateral gastrocnemius primarily involved in mid and terminal stance) and Module 3 (related to tibialis anterior and rectus femoris primarily involved in early stance, and early and late swing) were comparable across all subjects regardless of synergies number. PwMS had shorter stride length, longer double support phase and push off deficit with respect to HS (p<0.05). The alterations of activation timing profiles of specific modules in PwMS were associated with their walking deficits (e.g. the reduction of Module 2 activation percentage index in terminal stance, PwMS 35.55±13.23 vs HS 50.51±9.13% p<0.05, and the push off deficit, PwMS 0.181±0.136 vs HS 0.291±0.062 W/Kg p<0.05). Conclusion During gait PwMS have synergies numbers similar to healthy persons. Their neurological deficit alters modular control through modifications of the timing activation profiles rather than module composition. These changes were associated with their main walking impairment, muscle weakness and prolonged double support.
Motor modules are neural entities hypothesized to be building blocks of movement construction. How motor modules are underpinned by neural circuits has remained obscured. As a first step towards dissecting these circuits, we optogenetically evoked motor outputs from the lumbosacral spinal cord of two strains of transgenic mice - the Chat, with channelrhodopsin (ChR2) expressed in motoneurons, and the Thy1, expressed in putatively excitatory neurons. Motor output was represented as a spatial field of isometric ankle force. We found that Thy1 force fields were more complex and diverse in structure than Chat fields: the Thy1 fields comprised mostly non-parallel vectors while the Chat fields, mostly parallel vectors. In both, most fields elicited by co-stimulation of two laser beams were well explained by linear combination of the separately-evoked fields. We interpreted the Thy1 force fields as representations of spinal motor modules. Our comparison of the Chat and Thy1 fields allowed us to conclude, with reasonable certainty, that the structure of neuromotor modules originates from excitatory spinal interneurons. Our results not only demonstrate, for the first time using optogenetics, how the spinal modules follow linearity in their combinations, but also provide a reference against which future optogenetic studies of modularity can be compared.
Locomotion is produced by a central pattern generator. Its spinal cord organization is generally considered to be distributed, with more rhythmogenic rostral lumbar segments. While this produces a rostrocaudally traveling wave in undulating species, this is not thought to occur in limbed vertebrates, with the exception of the interneuronal traveling wave demonstrated in fictive cat scratching (Cuellar et al. J Neurosci 29:798–810, 2009). Here, we reexamine this hypothesis in the frog, using the seven muscle synergies A to G previously identified with intraspinal NMDA (Saltiel et al. J Neurophysiol 85:605–619, 2001). We find that locomotion consists of a sequence of synergy activations (A–B–G–A–F–E–G). The same sequence is observed when focal NMDA iontophoresis in the spinal cord elicits a caudal extension-lateral force-flexion cycle (flexion onset without the C synergy). Examining the early NMDA-evoked motor output at 110 sites reveals a rostrocaudal topographic organization of synergy encoding by the lumbar cord. Each synergy is preferentially activated from distinct regions, which may be multiple, and partially overlap between different synergies. Comparing the sequence of synergy activation in locomotion with their spinal cord topography suggests that the locomotor output is achieved by a rostrocaudally traveling wave of activation in the swing–stance cycle. A two-layer circuitry model, based on this topography and a traveling wave reproduces this output and explores its possible modifications under different afferent inputs. Our results and simulations suggest that a rostrocaudally traveling wave of excitation takes advantage of the topography of interneuronal regions encoding synergies, to activate them in the proper sequence for locomotion.
•Studies of modularity and compositionality provide insights into cortical organization.•Neural population activities are analyzed from representational and dynamic approaches.•Motor mapping and neural decoding studies promise new insights into motor organization.•New computational tools and theoretical modeling offer promise for further progress.
In this article we explore the complexities of what goes on in the brain when one wishes to perform even the simplest everyday movements. In doing so, we describe experiments indicating that the spinal cord interneurons are organized in functional modules and that each module activates a distinct set of muscles. Through these modules the central nervous system has found a simple solution to controlling the large number of muscle fibers active even during the execution of the simplest action. We also explore the many different neural signals that contribute to pattern formations, including afferent information from the limbs and information of motor memories.
Evidence suggests that the CNS uses motor primitives to simplify movement control, but whether it actually stores primitives instead of computing solutions on the fly to satisfy task demands is a controversial and still-unanswered possibility. Also in contention is whether these primitives take the form of time-invariant muscle coactivations ("spatial" synergies) or time-varying muscle commands ("spatiotemporal" synergies). Here, we examined forelimb muscle patterns and motor cortical spiking data in rhesus macaques (Macaca mulatta) handling objects of variable shape and size. From these data, we extracted both spatiotemporal and spatial synergies using non-negative decomposition. Each spatiotemporal synergy represents a sequence of muscular or neural activations that appeared to recur frequently during the animals' behavior. Key features of the spatiotemporal synergies (including their dimensionality, timing, and amplitude modulation) were independently observed in the muscular and neural data. In addition, both at the muscular and neural levels, these spatiotemporal synergies could be readily reconstructed as sequential activations of spatial synergies (a subset of those extracted independently from the task data), suggestive of a hierarchical relationship between the two levels of synergies. The possibility that motor cortex may execute even complex skill using spatiotemporal synergies has novel implications for the design of neuroprosthetic devices, which could gain computational efficiency by adopting the discrete and low-dimensional control that these primitives imply.SIGNIFICANCE STATEMENT:We studied the motor cortical and forearm muscular activity of rhesus macaques (Macaca mulatta) as they reached, grasped, and carried objects of varied shape and size. We applied non-negative matrix factorization separately to the cortical and muscular data to reduce their dimensionality to a smaller set of time-varying "spatiotemporal" synergies. Each synergy represents a sequence of cortical or muscular activity that recurred frequently during the animals' behavior. Salient features of the synergies (including their dimensionality, timing, and amplitude modulation) were observed at both the cortical and muscular levels. The possibility that the brain may execute even complex behaviors using spatiotemporal synergies has implications for neuroprosthetic algorithm design, which could become more computationally efficient by adopting the discrete and low-dimensional control that they afford.
AMERICAN SOCIETY OF PLASTIC SURGEONS PLASTIC & RECONSTRUCTIVE SURGERY PRS GLOBAL OPEN ASPS EDUCATION NETWORK AMERICAN SOCIETY OF PLASTIC SURGEONS PLASTIC & RECONSTRUCTIVE SURGERY PRS GLOBAL OPEN ASPS EDUCATION NETWORK
On page 6594, polymer fibers are designed by P. Anikeeva and co-workers to mimic fibrous geometry of the nerves allow simultaneous optical stimulation and neural recording in the spinal cord. The cover picture illustrates the flexibility of the bifunctional neural probes and their ability to accommodate deformation due to the movement of vertebrae.
Electrical microstimulation studies provide some of the most direct evidence for the neural representation of muscle synergies. These synergies, i.e., coordinated activations of groups of muscles, have been proposed as building blocks for the construction of motor behaviors by the nervous system. Intraspinal or intracortical microstimulation (ICMS) has been shown to evoke muscle patterns that can be resolved into a small set of synergies similar to those seen in natural behavior. However, questions remain about the validity of microstimulation as a probe of neural function, particularly given the relatively long trains of supratheshold stimuli used in these studies. Here, we examined whether muscle synergies evoked during ICMS in two rhesus macaques were similarly encoded by nearby motor cortical units during a purely voluntary behavior involving object reach, grasp, and carry movements. At each microstimulation site we identified the synergy most strongly evoked among those extracted from muscle patterns evoked over all microstimulation sites. For each cortical unit recorded at the same microstimulation site, we then identified the synergy most strongly encoded among those extracted from muscle patterns recorded during the voluntary behavior. We found that the synergy most strongly evoked at an ICMS site matched the synergy most strongly encoded by proximal units more often than expected by chance. These results suggest a common neural substrate for microstimulation-evoked motor responses and for the generation of muscle patterns during natural behaviors.
Inhibitory neurons in the adult mammalian spinal cord are known to locally modulate afferent feedback - from muscle proprioceptors and from skin receptors - to pattern motor activity for locomotion and postural control. Here, using optogenetic tools, we explored how the same population of inhibitory interneurons globally affects hindlimb movements in the spinal cord of both anesthetized and freely moving mice. Activation of inhibitory interneurons up to the middle/lower spinal cord i.e. T8-T9, were able to completely and globally suppress all ipsilateral hindlimb movements. Furthermore, the same population of interneurons - which inhibited movements - did not significantly change the sensory and proprioceptive information from the affected limbs to the cortex. These results suggest a rostro-caudal organization of inhibition in the spinal cord motor output without modulation of ascending sensory pathways.