Motor control
Motor control is the regulation of movements in organisms that possess a nervous system. Motor control includes conscious voluntary movements, subconscious muscle memory and involuntary reflexes, as well as instinctual taxes.
To control movement, the nervous system must integrate multimodal sensory information and elicit the necessary signals to recruit muscles to carry out a goal. This pathway spans many disciplines, including multisensory integration, signal processing, coordination, biomechanics, and cognition, and the computational challenges are often discussed under the term sensorimotor control. Successful motor control is crucial to interacting with the world to carry out goals as well as for posture, balance, and stability.
Some researchers argue that motor control is the reason brains exist at all.
Neural control of muscle force
All movements, e.g. touching your nose, require motor neurons to fire action potentials that results in contraction of muscles. In humans, ~150,000 motor neurons control the contraction of ~600 muscles. To produce movements, a subset of 600 muscles must contract in a temporally precise pattern to produce the right force at the right time.Motor units and force production
A single motor neuron and the muscle fibers it innervates are called a motor unit. For example, the rectus femoris contains approximately 1 million muscle fibers, which are controlled by around 1000 motor neurons. Activity in the motor neuron causes contraction in all of the innervated muscle fibers so that they function as a unit. Increasing action potential frequency in the motor neuron increases the muscle fiber contraction force, up to the maximal force. The maximal force depends on the contractile properties of the muscle fibers. Within a motor unit, all the muscle fibers are of the same type or Type II fibers ), and motor units of multiple types make up a given muscle. Motor units of a given muscle are collectively referred to as a motor pool.The force produced in a given muscle thus depends on: 1) How many motor neurons are active, and their spike rates; 2) the contractile properties and number of muscle fibers innervated by the active neurons. To generate more force, increase the spike rates of active motor neurons and/or recruiting more and stronger motor units. In turn, how the muscle force produces limb movement depends on the limb biomechanics, e.g. where the tendon and muscle originate and where the muscle inserts on the bone that it moves.
Recruitment order
Motor units within a motor pool are recruited in a stereotypical order, from motor units that produce small amounts of force per spike, to those producing the largest force per spike. The gradient of motor unit force is correlated with a gradient in motor neuron soma size and motor neuron electrical excitability. This relationship was described by Elwood Henneman and is known as Henneman's size principle, a fundamental discovery of neuroscience and an organizing principle of motor control.For tasks requiring small forces, such as continual adjustment of posture, motor units with fewer muscle fibers that are slowly-contracting, but less fatigueable, are used. As more force is required, motor units with fast twitch, fast-fatigeable muscle fibers are recruited.
High|
| _________________
Force required | /
| |
| |
| _____________|_________________
| __________|_______________________________
Low|__________|__________________________________________
↑ ↑ ↑ Time
Type I Recruit first Type II A Type IIB
Computational issues of motor control
The nervous system produces movement by selecting which motor neurons are activated, and when. The finding that a recruitment order exists within a motor pool is thought to reflect a simplification of the problem: if a particular muscle should produce a particular force, then activate the motor pool along its recruitment hierarchy until that force is produced.But then how to choose what force to produce in each muscle? The nervous system faces the following issues in solving this problem.
- Redundancy. Infinite trajectories of movements can accomplish a goal. How is a trajectory chosen? Which trajectory is best?
- Noise. Noise is defined as small fluctuations that are unrelated to a signal, which can occur in neurons and synaptic connections at any point from sensation to muscle contraction.
- Delays. Motor neuron activity precedes muscle contraction, which precedes the movement. Sensory signals also reflect events that have already occurred. Such delays affect the choice of motor program.
- Uncertainty. Uncertainty arises because of neural noise, but also because inferences about the state of the world may not be correct.
- Nonstationarity. Even as a movement is being executed, the state of the world changes, even through such simple effects as reactive forces on the rest of the body, causing translation of a joint while it is actuated.
- Nonlinearity. The effects of neural activity and muscle contraction are highly non-linear, which the nervous system must account for when predicting the consequences of a pattern of motor neuron activity.
"Optimal feedback control" is an influential theoretical framing of these computation issues.
Model systems for motor control
All organisms face the computational challenges above, so neural circuits for motor control have been studied in humans, monkeys, horses, cats, mice, fish lamprey, flies, locusts, and nematodes, among many others. Mammalian model systems like mice and monkeys offer the most straightforward comparative models for human health and disease. They are widely used to study the role of higher brain regions common to vertebrates, including the cerebral cortex, thalamus, basal ganglia and deep brain medullary and reticular circuits for motor control. The genetics and neurophysiology of motor circuits in the spine have also been studied in mammalian model organisms, but protective vertebrae make it difficult to study the functional role of spinal circuits in behaving animals. Here, larval and adult fish have been useful in discovering the functional logic of the local spinal circuits that coordinate motor neuron activity. Invertebrate model organisms do not have the same brain regions as vertebrates, but their brains must solve similar computational issues and thus are thought to have brain regions homologous to those involved in motor control in the vertebrate nervous system, The organization of arthropod nervous systems into ganglia that control each leg as allowed researchers to record from neurons dedicated to moving a specific leg during behavior.Model systems have also demonstrated the role of central pattern generators in driving rhythmic movements. A central pattern generator is a neural network that can generate rhythmic activity in the absence of an external control signal, such as a signal descending from the brain or feedback signals from sensors in the limbs. Evidence suggests that real CPGs exist in several key motor control regions, such as the stomachs of arthropods or the pre-Boetzinger complex that control breathing in humans. Furthermore, as a theoretical concept, CPGs have been useful to frame the possible role of sensory feedback in motor control.