The Organization of Movement From Ch. 33 “Principles of Neural Science”, 4th Ed. Kandel et al The sensory and motor systems • The sensory system integrates sensory information to construct internal representations – Physical energy is transformed into neural signals • The motor system uses internal motor representation to generate movement - Neural signals are transformed into physical energy • Purposeful action is only possible because the motor system has access to an on-going stream of sensory information The motor system • The motor system plans, coordinates and execute movements • The motor system is hierarchically organized – Spinal cord – Brain stem – Motor cortex • Successively higher levels of the motor hierarchy specify increasingly complex aspects of a motor task – Reflexive movements (withdrawal reflexes) – Rhythmic movements (chewing, swallowing, scratching) – Voluntary movements (goal directed) Reflexes • Reflexes are involuntary coordinated patterns of muscle contractions and relaxations elicited by peripheral stimuli – – • Different reflexes have different spatial and temporal patterns of muscle contractions Reflexes varies with stimulus site and receptor type activated Standardized response – – Same response time after time Modulation my context of behaviour by descending input • Reflexes circuits are used to coordinate muscles in complex voluntary movements • Reflexes and rhythmic motor patterns lie in the spinal cord and the brain stem Voluntary movements • Initiated to accomplish a specific goal • Improve with practice – Learning to anticipate and correct for obstacles that perturb the body • Take external perturbations into account – Feedback control: sensory feedback directly from the moving limb – Feed-forward control: use of multimodal sensory information to detect imminent perturbations to initiate proactive strategies (based on experience) Motor control circuits • Signals from sensors are compared with a desired state (reference signal) • The difference (error signal) is used to adjust the output • Feedback is especially important to maintain limb position or forces applied to objects • Without feedback the limb begins to drift as muscle fiber fatigue goes undetected Motor control circuits Example: Catching a ball • Feed-forward systems act in advance of certain perturbations • Experience is important to generate relevant anticipatory commands • Accuracy requires prior knowledge of trajectories • Without feedback the limb begins to drift as muscle fiber fatigue goes undetected Feed-forward: catching a ball 3 key principles for feed-forward control: Feed-forward is essential for rapid action Feed-forward depends on predicting the consequences of sensory events Feed-forward mechanisms can modify feedback mechanisms in the spinal cord Arrows: anticipatory response onset Modification of stretch reflex: flexor (F) and extensor (E) co-activated to stiffen elbow. Only spinal reflexes can mediate such rapid feedback Voluntary movements • 3 laws govern voluntary movements (modifiable by learning) 1. The brain represents the outcome of actions independently of the specific muscle used or the specific way the action is achieved 2. The time taken to respond to a stimulus depends on the amount of information needed to be processed 3. There is a trade-off between the speed of movement and the accuracy of the movement Voluntary movements • 3 laws govern voluntary movements (modifiable by learning) 1. The brain represents the outcome of actions independently of the specific muscle used or the specific way the action is achieved 2. The time taken to respond to a stimulus depends on the amount of information needed to be processed 3. There is a trade-off between the speed of movement and the accuracy of the movement T5 T6 Invariant features • • • All paths have similar shape All speed profiles have similar shapes Speed profiles scale linearly with the distance covered Variant features • Profiles of shoulder and elbow angles changes Conclusions • • Planning of movement is done with reference to the hand Represented by in abstract form rather than relative to joints or muscles Is the movement planned entirely before execution or is it continuously assesses? Is the movement planned? Speed and acceleration are scaled proportionately to the distance of the target The extent of movement is planned before the movement is initiated The representation of this plan is called a motor program Movement kinematics: spatial features and angles Distances: 2.5, 5, 10, 20, and 30 cm Movement dynamics: forces and torques Lifting a slippery object (Absence of feedback) (Constant force) Both feed-forward and feedback controls are used. (B) Load and grip force increase proportionally to the weight of the objects (scaling of preprogrammed force profile) (C) Training: 400 g object presented several times (dashed line). 800 g object then given. Force is increased in absence of sensory feedback via spinal circuit Voluntary movements • 3 laws govern voluntary movements (modifiable by learning) 1. The brain represents the outcome of actions independently of the specific muscle used or the specific way the action is achieved 2. The time taken to respond to a stimulus depends on the amount of information needed to be processed 3. There is a trade-off between the speed of movement and the accuracy of the movement Reaction time RT increases with choice and decreases with learning Reaction time indicates the amount of neural processing taking place (Also depends on sensory modality and conduction distance) Parallel processing of movement To overcome the slowness of serial processing, multiple stimuli and responses can be processed in parallel pathways >200 ms: trajectories are stereotyped (pre-programmed), match the target force and direction 0-100 ms: before extent or direction of movement is processed: no scaling, random direction 100-200 ms: Scaling but no direction = they are specified independently in parellel. Paradigm: Fixed arm, produce a pulse of elbow force (isometric) Match the peak of the force 6 possible target levels: three up (flexion) and three down (extension) Voluntary movements • 3 laws govern voluntary movements (modifiable by learning) 1. The brain represents the outcome of actions independently of the specific muscle used or the specific way the action is achieved 2. The time taken to respond to a stimulus depends on the amount of information needed to be processed 3. There is a trade-off between the speed of movement and the accuracy of the movement Accuracy and speed Fast movements are less accurate than slow ones due to less time for feedback corrections. A rapid increase in force requires recruitment of additional motor neurons A constant incremental increase in force is produced my a progressively smaller number of motor neurons As forces increases, fluctuations in the smaller number of motor neurons will be proportionately greater Learning will eliminate uncertainties and increase accuracy and speed Accuracy and learning Movement path becomes straighter and less variable Hierarchical organization • • The motor system can perform many different motor tasks with speed and accuracy – Sensory inputs to motor neurons and muscles is distributed in hierarchically interconnected areas of the spinal cord, brain stem and forebrain – Movement-related sensory information is processed in a different system that operates in parallel to the motor system – Reflex circuits in spinal cord and brain stem simplifies the instructions the cortex sends to lower levels 3 levels of organization – The spinal cord – The brain stem (medial/ lateral systems) – The cerebral cortex (direct/indirect projections) Hierarchical organization • • The cerebellum and basal ganglia – Feedback circuits for modulation – Forms two loop – Relay through different parts of thalamus – No direct output to the spinal cord – Are necessary for smooth movement and posture Diseases of the cerebellum and basal ganglia – Parkinson and Huntington diseases produce involuntary movements and abnormal posture – Cerebellar ataxias: abnormal timing and coordination of movements in progress The spinal cord Primary afferents from peripheral tissue • Connects to local interneurons • Connects to propriospinal interneurons • Connects to projection neurons to the brain • Connects to motor neurons The proximal-distal rule • Motor neurons innervating the most proximal muscles lie most medially • Motor neurons innervating distal muscles are located more laterally Propriospinal interneurons • • Short range projects dorsolaterally (distal limbs) Long range projects ventromedially (axial limbs) Hierarchical organization • The brain stem • • – Contains motor neurons controlling facial muscles – Projection neurons to the spinal cord – Receives descending input from the cerebral cortex – Other brain stem systems control movements of eyes and head Projection systems – The medial system: control of posture; integrates visual, vestibular and somatosensory input – The lateral systems: controls distal limb muscles; goal-directed movements of hand and arm Medial pathways – Vestibulospinal – Reticulospinal – Tectospinal tracts Ventromedial Dorsolateral