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    Psychology 110: Biological Psychology

    Lecture 13: Motor control

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    Outline

    we will look at the basic components of the motor system: muscles,motoneurons and proprioceptive receptors.

    we will then look at the hierarchy of processing that exists in the motorsystem, beginning with spinal reflexes, and continuing to look at the

    primary motor cortex, and motor association cortices.

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    Muscles

    Muscles can exert force only by contraction. The movement produced bythis contraction is determined by the muscles points of attachment(tendons).

    E.g. because of their attachments,contraction of the biceps flexes thearm, while contraction of the tricepsextends the arm (we call this

    muscular synergism).

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    Muscle fibers

    Muscle fibers consist of many individual fibers bound together withconnective tissue.

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    Mechanism of muscle contraction

    Muscle fibers consist of two types of interlocking protein actin and myosin.

    During contraction the myosin heads bind to actin, and then bend to slide

    filaments past one another, shortening the muscle.

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    Types of muscle fiber

    i). Fast muscle:

    reacts quickly, fatigues rapidly used for activities in which muscle tension changes frequently e.g.

    locomotion white meat in poultry (used to beat wings)

    ii). Slow muscle:

    react slowly but resist fatigue

    used to maintain posture dark meat in poultry (used for standing)

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    Innervation of muscle fiber

    Motoneurons leave the ventral root of the spinal cord and branch multipletimes before synapsing with individual muscle fibers at the neuromuscular

    junction.

    ACh is released and elicits anaction potential in the musclefiber itself.

    The influx of Ca2+ and Na+into the fiber triggers themolecular changes in actionand myosin that produce

    contraction.

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    Innervation ratio

    The innervation ratio is the ratio of motor axons to muscle fibers.

    Lower innervation ratios permit more fine motor control.

    For example, the muscles involved in eye movements have about one

    motoneuron for every three muscle fibers (1:3 innervation ratio), whilethose for the leg have one motoneuron controlling several hundred fibers(~1:300).

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    Motoneuron recruitment

    Motoneurons differ in size and electrophysiological properties.

    Large motoneurons have wide diameter axons to conduct impulses fast, butare difficult to initially excite.

    Small motoneurons have small axons and conduct impulses slowly, but areeasy to initially excite.

    SIZE PRINCIPLE:

    Muscle tension is increased by recruitment of

    increasing numbers of motoneurons in fixedorder according to their size. Weak stimulationactivates only the small neurons, and strongerstimulation recruits additional larger neurons.

    The principle is similar to range fractionation.Henneman

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    Sensory feedback (proprioception)

    There are two proprioceptive sensory structures that are associated withmuscles:

    i). Muscle spindles

    ii). Golgi tendon organs

    Proprioception refers to the detection of the position and movement of ourbody.

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    Muscle spindles

    A muscle spindle consists of a nerve fiber wrapped around an intrafusalmuscle fiber (extrafusal muscle fibers are responsible for the maincontraction of the muscle).

    If the muscle fiber is stretched this stretches the nerve fiber and causesthe nerve to fire action potentials.

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    Spinal reflexes: the stretch reflex

    1. Weight is placed insubjects hand.

    2. MA is stretched.

    3. SNA is excited.

    4. SNA connectsmonosynaptically withMNA exciting thestretched muscle.

    5. Via an interneuron,SNA also inhibits MNBensuring that the

    antagonistic muscle(MB) is relaxed whenMA is contracted.

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    Muscle spindles: efferent nerves

    Muscle spindles also receive input from efferent nerves called gamma

    motoneurons (alpha motoneurons go to main muscle fibers).

    Activity in gamma fibers causes a contraction in the length of the spindle,

    which modifies its sensitivity.

    The importance of this system is reflected in the fact that 30% of all

    motoneurons are gamma.

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    Golgi tendon organ

    Golgi tendon organs detect stretch in the tendon, not the muscle, and aresensitive to contraction.

    The Golgi tendon organ has feedback inhibition on the motoneuronsstimulating the muscle. If they detect a muscle overload (too muchcontraction) they fire, and inhibit the neuron that is causing the musclecontraction.

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    Outline

    we will begin by looking at the basic components of the motor system,including muscles, motoneurons and proprioceptive receptors.

    we will then look at the hierarchy of processing that exists in the motorsystem, beginning with spinal reflexes, and continuing to look at theprimary motor cortex, and motor association cortices (premotor cortexand the supplementary motor area.

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    Spinal reflexes

    An animal with damage to its spinal cord is still capable of many complexbehaviors. For example, the animal can often stand, and if stimulated, walkas well.

    Spinal cord contains a simple oscillator circuit, called a central patterngenerator, that can produce simple locomotion.

    At first cell A transmits thesame number of spikes ascell B

    Negative feedback reaches Bout of phase with the inputbecause of synaptic delays,

    causing rhythmic output.

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    Corticospinal pathway

    Consists of motoneurons that travel from the motor cortex, through thebrainstem, and down the spinal cord.

    Cross over in the medulla so that the left motor cortex controls the right sideof the body and vice versa.

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    Primary motor cortex (M1)

    The hands, lips and tongue are over represented in M1.

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    Spot the difference?

    SENSORY HOMUNCULUS MOTOR HOMUNCULUS

    In general, areas with many motor outputs also receive a lot of

    somatosensory input. There are some subtle differences, e.g. human earsreceive little motor output, and our eyes have few somatosensory inputs.

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    Primary motor cortex (M1)

    lesions of M1 cause paralysis

    electrical stimulation elicits twitches and jerks in limbs

    M1 neurons show direction selectivity for movements

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    Muscles or movements?

    Monkey must move joystick from central location to one of eight peripherallocations (indicate by yellow dot), but by altering the monkeys handorientation, one can alter which muscles are used to make the movement.

    One third of M1 neurons encode which muscles are used, one half encode themovement direction irrespective of the muscles used.

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    Premotor cortices

    Supplementary motor area (SMA) a.k.a. medial premotor cortex (MPC)

    Premotor cortex a.k.a. lateral premotor cortex (LPC)

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    LPC: involvement in conditional learning

    Conditional tasks involve learning an arbitrary relationship (If X then Y).

    TASK 1: Color instructs either pull or turna lever.

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    LPC: involvement in conditional learning

    Conditional tasks involve learning an arbitrary relationship (If X then Y).

    TASK 1: Color instructs either pull or turna lever.

    TASK 2: Color instructs which shape tochoose.

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    LPC: involvement in conditional learning

    (UC = unoperated control)

    Lesions of LPC impair conditional tasks, butonly when the stimulus instructs a specificmotor act.

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    Monkeys were trained to raise theirarms for a peanut. Monkeys with MPClesions were poor at doing this, failing toraise their arms as many times perminute as unoperated controls.

    Medial premotor cortex (MPC)

    NO CUE CUED

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    N i i l

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    Neuroimaging results

    simple movements activate M1

    increasing the motor complexity of the task extends the area of

    increased blood flow to include MPC

    when the sequence is indicated by external cues, lateral premotor cortex

    is activated instead

    when the subject mentally rehearses the complex movement, the area of

    activation is restricted to MPC

    M i i f i l hi h

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    Motor system organization: functional hierarchy

    PREFRONTAL

    Rules, strategies

    MEDIAL PREMOTOR

    Internally generatedactions

    LATERAL PREMOTOR

    Externally cued actions

    PRIMARY MOTOR

    Movements, muscles

    B i hi i t f

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    Brain-machine interface

    B i hi i t f (BMI)

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    Brain-machine interface (BMI)

    Matt Nagel was stabbed in the neckwhile breaking up a fight at a fireworksdisplay. Paralyzed from the neck down,he was implanted with an electrode

    array in his motor cortex (similar to themonkeys).

    With this array he was able to control a

    cursor on a computer screen (here heuses it to draw a circle) and type. Hewas also able to check email, surf theinternet and play some video games.

    Future hopes are that such devices willbe able to control prosthetic and roboticarms and control wheelchair. A further

    aim is to make the implants wireless.

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