Motor programs and central pattern generators

A motor program (MP) is a learned behavioral pattern defined as a neural network that can produce rhythmic output patterns with or without sensory input or central control. MPs are sets of movement commands, or “rules,” that define the details of skilled motor actions. An MP defines the specific muscles that are needed; the order of muscle activation; and the force, timing, sequence, and duration of muscle contractions. MPs help to control the degrees of freedom of interacting body structures and the number of ways in which each individual component acts. A generalized motor program (GMP) defines a pattern of movement with a lesser degree of specificity, rather than defining every individual aspect of a movement. GMPs allow for the adjustment, flexibility, and adaptation of movement features according to environmental demands. The existence of MPs and GMPs is a generally accepted concept; however, hard evidence that an MP or a GMP exists has yet to be found. Advancements in brain imaging techniques may substantiate this theory in the future. ^,

In contrast to MPs, a central pattern generator (CPG) is a genetically predetermined movement pattern. ^, CPGs exist as neural networks within the CNS and have the capability of producing rhythmic, patterned outputs resembling normal movement. These movements have the capability of occurring without sensory feedback inputs or descending motor inputs. Two characteristic signs of CPGs are that they result in the repetition of movements in a rhythmic manner and that the system returns to its starting condition when the process ceases. Both MPs and CPGs contribute to the development, refinement, production, and recovery of motor control throughout life.

The person, the task, and the environment: An ecological model for motor control

Motor control evolves so that people can cope with their environment. A person must focus on detecting information in the immediate environment (perception) that is determined to be necessary for performance of the task and achievement of the desired outcome goal. The individual is an active observer and explorer of the environment, which enables the development of multiple ways in which to accomplish (choose and execute) any given task. The individual analyzes a particular sensory environment and chooses the most suitable and efficient way to complete the task. The person consists of all functional and dysfunctional body structures and functions that exist and interact with one another. The task is the goal-directed behavior, challenge, or problem to be solved. The environment consists of everything outside of the body that exists, or is perceived to exist, in the external world. All three of these motor control constructs (person, task, environment) are dynamic and variable, and they interact with one another during learning and the production of a goal-directed, effective motor plan.

Role of sensory information in motor control

Sensory receptors from somatosensory (exteroceptors and proprioceptors), visual, and vestibular systems as well as taste, smell, and hearing fire in response to interactions with the external environment and to movement created by the body. Information about these various modalities is transmitted along afferent peripheral nerves to cells in the spinal cord and brain stem of the CNS. All sensory tracts with the exception of smell then synapse in respective sensory nuclei of the thalamus, which acts as a filter and relays this information to the appropriate lobe of the cerebral cortex (e.g., somatosensory to the parietal lobe, visual to the occipital lobe, vestibular, hearing, and taste to the temporal lobe). Sensory information is first received and perceived and then associated with other sensory modalities as well as memory in the association cortex. Once multiple sensory inputs are associated with one another, the person is able to perceive the body, its posture and movement, the environment and its challenges, and the interaction and position of the body with objects within the environment. The person uses this perceptual information to create an internal representation of the body (internal model) and to choose a movement program, driven by motivation and desire, to meet a final outcome goal. Although the sensory input and motor output systems operate differently, they are inseparable in function within the healthy nervous system. Agility, dexterity, and the ability to produce movement plans that are adaptable to environmental demands reflect the accuracy, flexibility, and plasticity of the sensorimotor system.

The CNS uses sensory information in a variety of ways to regulate posture and movement. Before movement is initiated, information about the position of the body in space, body parts in relation to one another, and environmental conditions is obtained from multiple sensory systems. Special senses of vision, vestibular inputs that respond to gravity and movement, and visual-vestibular interactions supply additional information necessary for static and dynamic balance and postural control as well as visual tracking. Auditory information is integrated with other sensory inputs and plays an important role in the timing of motor responses with environmental signals, reaction time, response latency, and comprehension of spoken word. This information is integrated and used in the selection and execution of a movement strategy. During movement performance, the cerebellum and other neural centers use feedback to compare the actual motor behavior with the intended motor plan. If the actual and intended motor behaviors do not match, an error signal is produced and alterations in the motor behavior are triggered. In some instances, the control system anticipates and makes corrective changes before the error signal is detected. This anticipatory correction is termed feedforward control. Changing one’s gait path while walking in a busy shopping mall to avoid a collision is an example of how visual information about the location of people and objects can be used in a feedforward manner.

Another role of sensory information is to revise the reference of correctness (central representation) of the MP before it is executed again. For example, a young child standing on a balance beam with the feet close together falls off of the beam. An error signal occurs because of the mismatch between the intended motor behavior and the actual motor result. If the child knows that his feet were too close together when he fell, then he will space his feet farther apart on the next trial. The child will then use this information about what happened, falling or not falling, in planning movement strategies for balancing on any narrow object such as a balance beam, log, or wall in the future.

Sensory information is necessary during the acquisition phase of learning a new motor skill and is useful for controlling movements during the execution of the motor plan. ^{, ,} However, sensory information is not always necessary when a person is performing well-learned motor behaviors in a stable and familiar context. ^, Rothwell and colleagues studied a man with severe sensory neuropathy in the upper extremity. He could write sentences with his eyes closed and drive a car with a manual transmission without watching the gear shift. He did, however, have difficulty with fine motor tasks such as buttoning his shirt and using a knife and fork to eat when denied visual information. The importance of sensory information must be weighed by the individual, who is unconsciously filtering and choosing appropriate and accurate sensory inputs to use to meet the movement goal.

Sensory experiences and learning alter sensory representations, or cortical “maps,” in the primary somatosensory, visual, and auditory areas of the brain. Training, as well as use and disuse of sensory information, has the potential to drive long-term structural changes in the CNS, including the formation, removal, and remodeling of synapses and dendritic connections in the cortex. This process of cortical plasticity is complex and involves multiple cellular and synaptic mechanisms. Plasticity in the nervous system is discussed further in the third section of this chapter.

Choice of motor pattern and the control of voluntary movement

A choice of body movement is made based on the person’s perception of the environment, his or her relationship to objects within it, and the goal to be met. The person chooses from a collection of plans that have been developed and refined over his or her lifetime. If a movement plan does not exist, a similar plan is chosen and modified to meet the needs of the task. Once the plan has been chosen, it is customized by the CNS with what are determined to be the correct actions to execute, given the perceived situation and goal of the individual.

Coordination

The movement plan is customized by communications between the frontal lobes, basal ganglia, and cerebellum, with functional connections through the brain stem and thalamus. During this process specific details of the plan are determined. Postural tone, coactivation, and timing of trunk muscle firing are set for proximal stability, balance, and postural control. Force, timing, and tone of limb synergies are set to allow for smooth, coordinated movements that are accurate in direction of trajectory, order, and sequence. The balance between agonist and antagonist muscle activity is determined so that fine distal movements are precise and skilled. This process is complicated by the number of possible combinations of musculoskeletal elements. The CNS must solve this “degrees of freedom” problem so that rapid execution of the goal-directed movement can proceed and reliably meet the desired outcome. ^, Once these movement details are complete, the motor plan is executed by the primary motor area in the precentral gyrus of the frontal lobe.

Execution of movement plans

Pyramidal cells in the corticospinal and corticobulbar tracts execute the voluntary motor plan. Neural impulses travel down these central efferent systems and communicate with motor neurons in the brain stem and spinal cord. The corticobulbar tract communicates with brain stem motor nuclei to control muscles of facial expression; mouth and tongue for speaking and eating; larynx and pharynx for voice and swallow; voluntary eye movements for visual tracking, saccades, and vergence; and muscles of the upper trapezius for shoulder girdle elevation. The corticospinal tract communicates with motor neurons in the spinal cord. The ventral corticospinal tract system communicates primarily with proximal muscle groups to provide the appropriate amount of voluntary activation needed to stabilize the trunk and limb girdles, thus allowing for dexterous distal limb movements. The lateral corticospinal tract system communicates primarily with muscles of the arms and legs—firing α motor neurons in coordinated synergy patterns with appropriate activity in agonist and antagonist muscles so that movements are smooth and precise. Other motor nuclei in the brain stem are programmed to fire just before corticospinal tract activity in order to supply postural tone and anticipatory muscle activity. These include lateral and medial vestibular spinal tracts, the reticulospinal tract, and the rubrospinal tract systems. Adequate and balanced muscle tone of flexors and extensors in the trunk and limbs is supplied automatically, without the need for conscious control. These brain stem nuclei have tonic firing rates that are modulated up or down to effectively provide more or less muscle tone in body areas depending on stimulation from gravity, limbic system activity, external perturbations, or other neuronal activity.

Adaptation

Adaptation is the process of using sensory inputs from multiple systems to adapt motor plans, decrease performance errors, and predict or estimate consequences of movement choices. The goal of adaptation is the production of consistently effective and efficient skilled motor actions. When all possible body systems and environmental conditions are considered in the motor control process, it is easy to understand why there is often a mismatch between the movement plan that is chosen and how it is actually executed. Errors in movements occur and cause problems that the nervous system must solve in order to deliver effective, efficient, accurate plans that meet the task goal. To solve this problem, the CNS creates an internal representation of the body and the surrounding world. This acts as a model that can be adapted and changed in the presence of varying environmental demands. It allows for the ability to predict and estimate the differences between similar situations. This ability is learned by practicing various task configurations in real-life environments. Without experience, accurate movement patterns that consistently meet desired task goals are difficult to achieve.

Anticipatory control

Anticipatory control of posture and postural adjustments stabilizes the body by minimizing displacement of the center of gravity. Anticipatory control involves motor plans that are programmed to act in advance of movement. A comparison between incoming sensory information and knowledge of prior movement successes and failures enables the system to choose the appropriate course of action. ^,

Flexibility

A person should have enough flexibility in performance to vary the details of a simple or complex motor plan to meet the challenge presented by any given environmental context. This is a beneficial characteristic of motor control. When considering postural control, for example, a person will typically display a random sway pattern during standing that may ensure continuous, dynamic sensory inputs to multiple sensory systems. The person is constantly adjusting posture and position to meet the demand of standing upright (earth vertical), as well as to seek information from the environment. Rhythmic, oscillating, or stereotypical sway patterns that are unidirectional in nature are not considered flexible and are not as readily adaptable to changes in the environment. Lack of flexibility in postural sway may actually render the person at greater risk for loss of balance and falls.

Role of the cerebellum

The primary roles of the cerebellum in motor control are to maintain posture and balance during static and dynamic tasks and to coordinate movements before execution and during performance. The cerebellum processes multiple neural signals from (1) motor areas of the cerebral cortex for motor planning, (2) sensory tract systems (dorsal spinal cerebellar tract, ventral spinal cerebellar tract) from muscle and joint receptors for proprioceptive and kinesthetic sense information resulting from movement performance, and (3) the vestibular system for the regulation of upright control and balance at rest and during movements. It compares motor plan signals driven by the cortex with what is received from muscles and joints in the periphery and makes necessary adjustments and adaptations to achieve the intended coordinated movement sequence. “Instructions” for movements that are frequently repeated are stored in the cerebellum as procedural memory traces. This increases the efficiency of its role in coordinating movement. The cerebellum also plays a role in the function of the reticular activating system (RAS). The RAS network exists in the brain stem tegmentum; it is a network of nerve cells that maintain human consciousness and help people focus attention and block out distractions that may affect motor performance. Damage to the cerebellum, its tract systems, or its structure creates problems of movement coordination—not execution or the choice of which program to run. The cerebellum also plays a role in language, attention, and mental imagery functions that are not considered to take place in motor areas of the cerebral cortex (see Table 3.2 ).

The cerebellum plays the following four important roles in motor control :

• 1.
  

  Feedforward processing : The cerebellum receives neural signals, processes them in a sequential order, and sends information out, providing a rapid response to any incoming information. It is not designed to act like the cerebral cortex and does not have the ability to generate self-sustaining neural patterns.

  
  
• 2.
  

  Divergence and convergence : The cerebellum receives a great number of inputs from multiple body structures. It processes this information extensively through a structured internal network and sends the results out through a limited number of output cells.

  
  
• 3.
  

  Modularity : The cerebellum is functionally divided into independent modules—hundreds to thousands—all with different inputs and outputs. Each module appears to function independently, although each shares neurons with the inferior olives, Purkinje cells, mossy and parallel fibers, and deep cerebellar nuclei.

  
  
• 4.
  

  Plasticity : Synapses within the cerebellar system (between parallel fibers and Purkinje cells and synapses between mossy fibers and deep nuclear cells) are susceptible to modification of their output strength. The influence of input on nuclear cells is adjustable, which provides great flexibility in adjusting and fine-tuning the relationship between cerebellar inputs and outputs.

Role of the basal ganglia

The basal ganglia are a collection of nuclei located in the forebrain and midbrain and consisting of the globus pallidus, putamen, caudate nucleus, substantia nigra, and subthalamic nuclei. The basal ganglia have primary functions in motor control and motor learning. They play a role in deciding which motor plan or behavior to execute at any given time and have connections to the limbic system; they are therefore believed to be involved in “reward learning.” The basal ganglia play a key role in eye movements through midbrain connections with the superior colliculus and help to regulate postural tone as a basis for the control of body positions, preparedness, and central set (i.e., the nervous system’s internal model of strategies for reactive balance control). Refer to Chapter 18 for additional information on the basal ganglia.

Movement patterns arising from self-organizing subsystems

Coordinated movement patterns are developed and refined via dynamic interaction among body systems and subsystems in response to internal and external constraints. Movement patterns used to accomplish a goal are contextually appropriate and arise as an emergent property of subsystem interaction. Several principles relate to self-organizing systems: reciprocity, distributed function, consensus, and emergent properties.

Reciprocity implies information flow between two or more neural networks. These networks can represent specific brain centers—for example, the cerebellum and basal ganglia ( Fig. 3.2 ). Alternatively, the neural networks can be interacting neuronal clusters located within a single center—for example, the basal ganglia. One model to demonstrate reciprocity is the basal ganglia regulation of motor behavior through direct and indirect pathways to cortical areas. The more direct pathway from the putamen to the globus pallidus internal segment provides net inhibitory effects. The more indirect pathway from the putamen through the globus pallidus external segment and subthalamic nucleus provides a net excitatory effect on the globus pallidus internal segment. Alteration of the balance between these pathways is postulated to produce motor dysfunction. ^, An abnormally decreased outflow from the basal ganglia is postulated to produce involuntary motor patterns; these produce excessive motion such as chorea, hemiballism, or nonintentional tremor. Alternatively, an abnormally increased outflow from the basal ganglia is postulated to produce a paucity of motions, as seen in the rigidity observed in individuals with Parkinson disease (see Chapter 18 ).

Systems Model of Motor Control.

Distributed function presupposes that a single center or neural network has more than one function. The concept also implies that several centers share the same function. For example, a center may serve as the coordinating unit of an activity in one task and as a pattern generator or oscillator to maintain the activity in another task. An advantage of distributing function among groups of neurons or centers is to provide centers with overlapping or redundant functions. Neuroscientists believe such redundancy is a safety feature. If a neuronal lesion occurs, other centers can assume critical functional roles, thereby producing recovery from CNS dysfunction.

Consensus implies that motor behavior occurs when a majority of brain centers or regions reach a critical threshold to produce activation. Also, through consensus, extraneous information or information that does require immediate attention is filtered. If, however, a novel stimulus enters the system, it carries more weight and receives immediate attention. A novel stimulus may be new to the system, may reflect a potentially harmful situation, or may result from the conflict of multiple inputs.

Emergent properties may be understood by the adage “the whole is greater than the sum of its parts.” This concept implies that brain centers, not a single brain center, work together to produce movement. An example of the emergent properties concept is continuous repetitive activity (oscillation). In Fig. 3.3 A, a hierarchy is represented by three neurons arranged in tandem. The last neuron ends on a responder. If a single stimulus activates this network, a single response occurs. What is the response if the neurons are arranged so that the third neuron sends a collateral branch to the first neuron in addition to the ending on the responder? In this case (see Fig. 3.3 B), a single stimulus activates neuron No. 1, which in turn activates neurons No. 2 and No. 3, causing a response as well as reactivating neuron No. 1. This neuronal arrangement produces a series of responses rather than a single response. This process is also termed endogenous activity.

Emergent Property.

Another example of an emergent property is the production of motor behavior. Rather than having every MP stored in the brain, an abstract representation of the intended goal is stored. At the time of motor performance, various brain centers use the present sensory information combined with past memory of the task to develop the appropriate motor strategy. This concept negates a hardwired MP concept. If MPs were hardwired and if an MP existed for every movement ever performed, the brain would need a huge storage capacity and would lack the adaptability necessary for complex function.

Controlling the degrees of freedom

Combinations of muscle and joint action permit a large number of degrees of freedom that contribute to movement. A system with a large number of degrees of freedom is called a high-dimensional system. For a contextually appropriate movement to occur, the number of degrees of freedom must be constrained. Bernstein suggested that the number of degrees of freedom could be reduced by muscles working in synergies—that is, coupling muscles and joints of a limb to produce functional patterns of movement. The functional unit of motor behavior is then a synergy. Synergies help to reduce the degrees of freedom, transforming a high-dimensional system into a low-dimensional system. For example, a step is considered to be a functional synergy pattern for the lower extremity. Linking together stepping synergies with the functional synergies of other limbs creates locomotion (interlimb coordination).

Functional synergy implies that muscles are activated in an appropriate sequence and with appropriate force, timing, and directional components. These components can be represented as fixed or “relative” ratios, and the control comes from input given to the cerebellum from higher centers in the brain and the peripheral or spinal system and from prior learning (see Chapter 19 ). ^{, ,} The relative parameters are also termed control parameters. Scaling control parameters leads to a change in motor behavior to accomplish the task. For example, writing your name on the blackboard exemplifies scaling force, timing, and amplitude. Scaling is the proportional increase or decrease of the parameter to produce the intended motor activity.

Coordinated movement is defined as an orderly sequence of muscle activity in a single functional synergy or the orderly sequence of functional synergies with appropriate scaling of activation parameters necessary to produce the intended motor behavior. Uncoordinated movement can occur at the level of the scaling of control parameters in one functional synergy or inappropriate coupling of functional synergies. The control parameter of duration will be used to illustrate scaling. If muscle A is active for 10% of the duration of the motor activity and muscle B is active 50% of the time, the fixed ratio of A/B is 1:5. If the movement is performed slowly, the relative time for the entire movement increases. Fixed ratios also increase proportionally. Writing your name on a blackboard very small or very large yields the same result—your name.

Timing of muscle on/off activation for antagonistic muscles such as biceps and triceps or hamstrings and quadriceps must be accurate for the coordination and control of movement patterns. If one muscle group demonstrates a delayed onset or maintains a longer duration of activity overlapping with triceps “on” time, the movement will appear uncoordinated. Patients with neurological dysfunction often demonstrate alterations in the timing of muscle activity within functional synergies and in coupling functional synergies to produce movement. ^, These functional movement synergies are not hardwired but represent emergent properties. They are flexible and adaptable to meet the challenges of the task and the environmental constraints.

Finite number of movement strategies

The concept of emergent properties could conceivably imply an unlimited number of movement strategies available to perform a particular task. However, limiting the degrees of freedom decreases the number of strategies available for selection. In addition, constraints imposed by the internal environment (e.g., musculoskeletal system, cardiovascular system, metabolic activity, cognition) and external environment (e.g., support surface, obstacles, lighting) limit the number of movement strategies. Horak and Nashner observed that a finite number of balance strategies were used by individuals in response to externally applied linear perturbations on a force plate system. With use of a life-span approach, VanSant identified a limited number of movement patterns for the upper limb, head-trunk, and lower limb for the task of rising from supine to standing.

The combination of these strategies produces the necessary variability in motor behavior. Although an individual has a preferred or modal profile, the healthy person with an intact neuromuscular system can combine strategies in various body regions to produce different movement patterns that also accomplish the task. ^, Persons with neurological deficits may be unable to produce a successful, efficient movement pattern because of their inability to combine strategies or adapt a strategy for a given environmental change (e.g., differing chair height for sit-to-stand transitions).

Variability of movements implies normalcy

A key to the assessment and treatment of individuals with neurological dysfunction lies in variability of movement and in the notion that variability is a sign of normalcy, and stereotypical behavior is a sign of dysfunction.

Age, activity level, the environment, constraints of a goal, and neuropathological conditions affect the selection of patterns available for use during movement tasks. When change occurs in one or more of the neural subsystems, a new movement pattern emerges. The element that causes change is called a control parameter. For example, an increase in the speed of walking occurs until a critical speed and degree of hip extension are reached, thereby switching the movement pattern to a run. When the speed of the run is decreased, there is a shift back to the preferred movement pattern of walking. A control parameter shifts the individual into a different pattern of motor behavior.

This concept underlies theories of development and learning. Development and learning can be viewed as moving the system from a stable state to a more unstable state. When the control variable is removed, the system moves back to the early, more stable state. As the control variable continues to push the system, the individual spends more time in the new state and less time in the earlier state until the individual spends most of the time in the new state. When this occurs, the new state becomes the preferred state. Moving or shifting to the new, preferred state does not obviate the ability of the individual to use the earlier state of motor behavior. Therefore new movement patterns arise when critical changes occur in the system because of a control parameter, but they do not eliminate older, less-preferred patterns of movement.

Motivation to accomplish a task in spite of functional limitations and neuropathological conditions can also shift the individual’s CNS to select different patterns of motor behavior. The musculoskeletal system, by nature of the architecture of the joints and muscle attachments, can be a constraint on the movement pattern. An individual with a functional contracture may be limited in the ability to bend a joint only into a desired range, thereby decreasing the movement repertoire available to him or her. Such a constraint produces adaptive motor behavior. Dorsiflexion of the foot needs to meet a critical degree of toe clearance during gait. If there is a range of motion limitation in dorsiflexion, then biomechanical constraints imposed on the nervous system will produce adaptive motor behaviors (e.g., to achieve toe clearance during gait). Changes in motor patterns during the task of rising from supine to standing are observed when healthy individuals wear orthoses to limit ankle dorsiflexion. The inability to easily open and close the hand with rotation may lead to adaptations that require the shoulder musculature to place the hand in a more functional position. This adaptation uses axial and trunk muscles and will limit the use of that limb in both fine and gross motor performance. Refer to Chapter 19 for more information in this regard.

Preferred, nonobligatory movement patterns that are stable yet flexible enough to meet ever-changing environmental conditions are considered attractor states. Individuals can choose from a variety of movement patterns to accomplish a given task. For example, older adults may choose from a variety of fall-prevention movement patterns when faced with the risk of falling. The choice of motor plan may be negatively influenced by age-related declines in the sensory input systems or a fear of falling. For example, when the Multi-Directional Reach Test is being performed, an older adult may choose to reach forward, backward (lean), or laterally without shifting her center of gravity toward the limits of stability. She has the capability of performing a different reaching pattern if asked but prefers a more stable pattern.

Obligatory and stereotypical movement patterns suggest that the individual does not have the capability of adapting to new situations or cannot use different movement patterns to accomplish a given task. This inability may be a result of internal constraints that are functional or pathophysiological. The patient who has had a stroke has CNS constraints that limit the number of different movement patterns that can emerge from the self-organizing system. With recovery, the patient may be able to select and use additional movement strategies. Cognition and the capability to learn may also limit the number of movement patterns available to the individual and the ability of the person to select and use new or different movement patterns.

Obligatory and stereotypical movement patterns also arise from external constraints imposed on the organism. Consider the external constraints placed on a concert violinist. External constraints can include, for example, the length of the bow and the position of the violin. Repetitive movement patterns leading to cumulative trauma disorder in healthy individuals can lead to muscular and neurological changes. Over time, changes in dystonic posturing and changes in the somatosensory cortex have been observed. Although one hypothesis considers that the focal dystonia results from sensory integrative problems, the observable result is a stereotypical motor problem.

To review, the nervous system responds to a variety of internal and external constraints to develop and execute motor behavior that is efficient to accomplish a specific task. Efficiency can be examined in terms of metabolic cost to the individual, type of movement pattern used, preferred or habitual movement (habit) used by the individual, and time to complete the task. The term attractor state is used in dynamical systems theory to describe the preferred pattern or habitual movement.

Individuals with neurological deficits may have limited repertoires of movement strategies available. Patients experiment with various motor patterns in order to learn the most efficient, energy-conscious motor strategy to accomplish the task. Therapists can plan interventions that help to facilitate refinement of the task to match the patient’s capability, allowing the task to be completed using a variety of movement strategies rather than limited stereotypical strategies, leading to improved function.

Errors in motor control

When the actual motor behavior does not match the intended motor plan, an error in motor control is detected by the CNS. Common examples of errors in motor control are loss of balance; inappropriate scaling of force, timing, or directional control; and inability to ignore unreliable sensory information, resulting in sensory conflict. Any one or combination of these errors may be the cause of a fall or an error in performance accuracy.

Errors also occur when unexpected factors disrupt the execution of the MP. For example, when the surface is unreliable (unstable, moving), this will force the individual to adapt motor responses to meet the demand of the environment. Switching between closed environments (more stable) and open environments (more unpredictable) will challenge the individual to adapt motor responses. When an individual steps off a moving sidewalk, a disruption in walking occurs. The first few steps are not smooth because the person has to switch his movement strategies from one incorporating a moving support surface to one incorporating a stationary support surface.

Errors occur in the perception of sensory information, in the selection of the appropriate MP, in the selection of the appropriate variable parameters, or in the response execution. Patients with neurological deficits may demonstrate a combination of these errors. Therefore an assessment of motor deficits includes analysis of these types of errors. If a therapist observes a motor control problem, there is no guarantee that the problem has arisen from within the motor system. Somatosensory problems can drive motor dysfunction; cognitive and emotional problems express themselves through motor output. Thus it is up to the movement specialist to differentiate the cause of the problem through valid and reliable examination tools (see Chapter 8 ). Once the cause of the motor problem has been identified, selection of interventions should lead to positive outcomes.

All individuals, both healthy and those with CNS dysfunction, make errors in motor programming. These errors are assessed by the CNS and are stored in past memory of the experience. Errors in motor programming are extremely useful in learning. Learning can be viewed as decreasing the mismatch between the intended and actual motor behavior. This mismatch is a measure of the error; therefore a decrease in the degree of the error is indicative of learning. Errors, then, are an important part of the rehabilitation process. However, this does not mean that the therapist allows the patient to practice errors over and over. The ability of the patient to detect an error and correct it to produce appropriate and efficient motor behavior is one key to recovery and an important consideration when intervention strategies are developed. This is discussed further in the next section of this chapter.

Variables related to the individual

The clinician must first differentiate general motor performance factors that are under the control of the individual’s cognitive and emotional systems and those that are controlled by the motor system itself. These concepts are presented in Fig. 3.5 . There are many cognitive factors—such as arousal, attention, and memory, as well as cortical pathways related to declarative or executive learning—that have specific influences over behaviors observed after neurological insult. ^, Other factors—such as limbic connections to cortical pathways affected by motivation, fear and belief, and emotional stability and instability—also dramatically affect motor performance and declarative learning. Some of these factors may also limit activity and participation. Therapists must learn how to discriminate between problems due to motor output, somatosensory input, cortical processing, and limbic emotional state problems and to identify how the last two systems affect motor output. With that differentiation, clinicians should also be able to separate specific motor system deficits from motor control problems arising from dysfunction within other areas of the CNS. Last, the patient’s fitness level; current limitations in strength, endurance, power, and range of motion; or pain level may also influence learning. ^,

Concepts Affecting Motor Learning.

Variables related to the task

The two major variables related to the task itself that must be considered when facilitating motor learning include practice of the task and feedback related to task performance.

Practice.

Practice is defined as “repeated performance in order to acquire a “skill.” As the definition implies, multiple repetitions of the task are usually required to be able to achieve skillful performance of a task. With other variables being constant, more practice results in more learning. To be effective, these repetitions must involve a process of problem solving rather than just repetition of the activity. The therapist can manipulate variables related to practice in order to optimize motor learning in an individual with a pathological movement pattern.

Practice conditions.

The term practice conditions refers to the manner in which the task or exercise is repeated with respect to rest periods, the amount of exercise, and the sequence in which these tasks or exercises are performed.

According to the apportionment of practice in relation to rest periods, massed tasks or exercises can be classified as massed practice or distributed practice. Massed practice is when the rest period is much shorter in relation to the amount of time the task or exercise is practiced. This is contrasted against distributed practice, in which the time between practice sets is equal to or greater than the amount of time devoted to practicing a particular task or activity such that the rest period is spread out throughout the practice. In terms of neurological physical therapy practice, it is important to consider the effect of physical and mental fatigue when a person is training. For example, physical fatigue sets in during massed practice of a particular balance exercise activity in standing and may cause a patient to fall. Moreover, individuals who are cognitively impaired may not respond positively to sustained activity that requires considerable concentration and therefore might fail in the performance of the skill. On the other hand, to be functional and useful in daily life, certain activities have to be performed without significant amounts of rest. For example, taking significant rest breaks when one is ambulating for even a short distance limits the person’s ability to use walking in a functional manner. Sometimes a patient needs more rest periods in the initial stages of learning a skill to compensate for impairments in muscular endurance or cardiopulmonary function with the intent of decreasing these rest periods to achieve skill performance that reflect how that activity is used in real-life situations. Therefore therapists should consider the skill demands and desired results when they are choosing one practice type versus another.

Complete, functional tasks or activities can usually be divided into smaller subcomponents for practice. The way those subcomponents are practiced relative to the entire task or activity can be manipulated to optimize motor learning. This includes practice of the entire task or parts of the task, whole learning, pure-part learning, progressive-part learning, or whole-part learning methods. Fig. 3.6 summarizes these concepts.

Type of Movement (Task) and Practice Environment.

Whole learning suggests that the learner practice the entire movement as one activity. Asking a person to stand up incorporates the entire activity of coming to stand from sitting. Simple movements such as rolling, coming to sit, coming to stand, and walking might best be taught as a whole activity as long as the individual has all the component parts to practice the whole.

In pure-part learning the therapist introduces one part first; then this part is practiced by the learner before another new part is introduced and practiced. Each part is critical to the whole movement, but which one is learned first does not matter. Learning a tennis serve is an excellent example of an activity that can be taught as a pure part. Learning to toss the ball vertically to a specific spot in space is a very different and separate part from swinging the tennis racket as part of the serve. Learning to squeeze the toothpaste onto the brush is a very different movement strategy from brushing the teeth.

Progressive-part learning is used when the sequence of movements and the component parts to be learned need to be programmed in a specific order. Line dancing is an activity taught using progressive part learning. Individuals with sequencing deficits must often be taught using progressive parts, or they will mix up the ordering of parts during an activity. Therapists see this in the clinical arena when an individual stands up from a wheelchair and then tries to pick up the foot pedals and lock the brakes. Given that problem, such a patient must practice progressive-part learning by first locking the brakes, then picking up the foot pedals, and finally standing. If the activity is not practiced using progressive-part learning, the patient will have little consistency in how the parts are put together, thereby placing that him at high risk of failing at the functional task.

Whole-part learning can be used when the skill or activity can be practiced between the whole and the parts. In the clinical environment, a common application of this concept is whole to part to whole learning . First the therapist has the patient try the whole activity, such as coming to stand or opening a door. Next, the therapist has the patient practice a component part that is deficient (e.g., gripping the doorknob). Finally the whole activity is practiced as a functional pattern. In this way therapists work on the functional activity and then work on correcting the impairment or limitation, such as power production, range, or balance. Then they go back to the functional activity in order to incorporate the part learning into the whole. An example might be asking a patient to first stand up from a chair. As she tries to stand she generates too much power, holds her breath, and cannot repeat the activity more than once. The therapist decides to practice a component part by first assisting the patient to a relaxed standing posture, then having her eccentrically begin to sit into a partial squat, and then having her return to standing. As the patient practices, she will increase the range of motion and eventually will sit and return to stand. Once that is accomplished, she will continue to practice sit to stand to sit as a whole activity.

Therapists change the sequence in which component parts of a task are practiced using blocked or random practice. In blocked practice the patient first practices a single task over and over before moving to the next task. With random practice the component skills are practiced in random order without any particular sequence. The contextual interference effect explains the difference in motor performance found when these two types of practice are compared. Studies have shown that performance may be enhanced by using blocked practice; however, learning is not enhanced by using this type of practice. Random practice has been shown to enhance learning because this type of practice forces the learners to come up with a motor solution each time a task is performed. ^,

Feedback.

The use of feedback is another important variable related to motor learning. Feedback is defined as the use of sensory information—visual, auditory, or somatosensory—to improve performance, retention, or transfer of a task. Internal feedback pertains to sensory information that the patient receives that can be used to improve performance of that particular task or activity in the future. The therapist provides extrinsic or augmented feedback with the intent of improving learning of the task. In people with neurological dysfunctions, extrinsic feedback is important because the patient’s intrinsic feedback system may be impaired or absent.

Extrinsic feedback can further be classified as knowledge of performance (KP) or knowledge of results (KR). KP is given concurrently while the task is being performed and can therefore also be called concurrent feedback. Feedback given concurrently, especially during the critical portions of the task, allows the patient to successfully perform the activity.

KR pertains to feedback given at the conclusion of the task (therefore it is also called terminal feedback ) and provides the patient with information about the success of his or her actions with respect to the activity. KR can be classified as faded, delayed, or summary. In faded feedback the therapist provides more information in the beginning stages of learning of the skill and slowly withdraws that feedback as the patient demonstrates improvement in the performance of the task. With delayed feedback, information is given to the patient when a period of time has elapsed after the task has been completed. The intent of this pause between the termination of a task and feedback is to give the patient some time to process the activity and generate possible solutions to the difficulties encountered in the previous performance of the task. In contrast, summary feedback is provided after the patient has performed several trials of a particular task without receiving feedback. Studies show that subjects who were given more frequent feedback, compared with those who received summary feedback, performed better during the task acquisition stage of learning but worse on retention tests. ^,

Additional concepts related to long-term learning are presented in Fig. 3.7 .

Concepts Important to Long-term Learning.

Variables related to the environment

Therapists can alter environmental conditions to optimize motor learning. Gentile ^, described the manipulation of the environment in which a task is performed to make an activity more appropriate for what the patient is able to do. Patients with neurological dysfunction may benefit from practicing in a closed environment that is stationary and predictable. It allows the skill to be practiced with minimal distractions or challenges from the environment, enabling the patient to plan the movement in advance. An example of this would be performing gait training in a quiet, empty therapy gym. As the patient improves, it is important to practice this activity in an open environment that is less predictable, more distracting, or in motion to provide a real-world application of the task. An example of this would be gait training the patient in a busy gym environment, in a crowded cafeteria, on a moving walkway, or in a noisy hallway.

If prior procedural learning has occurred, then creating an environment that allows the program to run in the least restrictive environment should lead to the most efficient outcome in the shortest time. ^, If a patient must learn a new program, such as walking with a stereotypical extension pattern, then goal-directed, attended practice with guided feedback will be necessary. It may be easier to bring back an old ambulatory pattern by creating an environment to elicit that program than to teach a patient to use a new inefficient movement program.

A therapist must identify what MPs are available and under what conditions. This allows the therapist to (1) determine whether deficits are present, (2) anticipate problems in performance, and (3) match existing programs with functional activity challenges during training. Similarly, knowing available MPs and the component body systems necessary to run those programs aids the therapist in the selection of intervention procedures.

If the patient has permanent damage to either the basal ganglia or the cerebellum, then retaining the memory of new MPs may be difficult and substitution approaches may become necessary. Through evaluation, the clinician must determine whether anatomical disease or a pathological condition is actually causing procedural learning problems and whether identifying and teaching a substitution pattern or teaching the patient to compensate with an old pattern will allow him to succeed at the task. However, therapists should never forget that the plasticity of the CNS can promote significant recovery and adaptation through the performance of attended, goal-directed, repetitive behavior. ^,

The provision of an appropriate level of challenge to the learner can optimize motor learning. The clinician must learn to expertly manipulate the environment to best facilitate learning. A task that is too difficult for the patient will result in persistent failure of performance, frustration, and lack of learning; then the only option will be to compensate through available patterns of movement that limit function. An activity that is too easy and routinely results in 100% success also does not result in learning because the learner becomes bored and no longer attends to the experience. The most beneficial level of challenge for training will create some errors in performance, require the patient to solve problems to meet the demands presented, and allow a level of success that inspires continued motivation to practice and achieve a higher standard of skill.