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MOTOR SYSTEMS: CONTROL OF MOVEMENT & BEHAVIOR

 
TABLE OF CONTENTS


Updated: Mar. 6, 2008


LECTURE INFORMATION

KEY CONCEPTS IN THIS LECTURE

1. The Autonomic Nervous System (ANS) and the Somatic Nervous System make up the efferent paths of the Peripheral Nervous System. The ANS is further subdivided into the sympathetic and parasynpathetic nervous systems, both of which are involved in integration of involuntary physiological processes, such as control of heart rate. Often, but not always, these two subsystems work in opposition to regulate a physiological event. The Somatic Nervous System controls either voluntary or involuntary motor events which always involve muscle contraction.

2. Nerves releasing Ach at the neuromuscular junction (=end plate) cause the contraction of skeletal muscle. The functional unit of a muscle organ is the muscle fiber (=muscle cell). The muscle fiber contracts in an "all-or-none" fashion when stimulated by an action potential. The action potential first causes intracellular Ca++ release from the sarcoplasmic reticulum and the Ca++ activates a cascade of events which results in the movement of actin over myosin (=sliding filament theory). The contractile process is achieved by the repeated formation of cross bridges between the myosin and actin myofibrils along with expenditure of ATP.

3. Muscles tend to maintain a status quo of not contracting until instructed to do otherwise. Some simple reflexes, such as the knee jerk reflex, illustrate this point. In this reflex, stretch of the muscle spindle in the muscle activates a neural circuit which causes contraction of extrafusal muscle. There are many reflexes, some of which have been incorporated into basic circuits associated with simple involuntary behaviors, such as walking in humans or swimming in the lamprey. Neural circuits which control such simple behaviors are often controlled by central pattern generators.

4. The brain controls most behavior. Sensory inputs from different cortical regions collect in the prefrontal cortex. Using this information, the secondary motor area (SMA) and the premotor area (PMA) organize the behavior and the primary motor cortex (M1) executes it. The descending paths include the lateral spinal columns which are important in hand and limb movements while the ventrolateral columns are involved with involuntary behaviors, such as walking, breathing, and eye movements. The basal ganglion also receive output from a number of cortical areas; these structures provide positive feedback to the SMA and PMA and may help in the initiation of a behavior. The cerebellum also receives cortical input and provides M1 with information on the status of a behavior as well as information on its feasibility.

LECTURE OBJECTIVES

1. The organization and function of the sympathetic and parasympathetic nervous systems will be discussed.
2. How a nerve innervates and activates the muscle cell
3. Establish the molecular basis for muscle contraction
4. Discuss how the CNS alters the status quo of a neural circuit to cause contraction of muscle
5. Establish how central pattern generators control basic rhythmic behaviors

LECTURE OUTLINE

I. INTRODUCTION **

  A. Review: The nervous system is made up of the CNS and PNS
     1. CNS: 
          a. Brain controls motor behavior
          b. Spinal cord relays information to and from the brain
               1) Ascending and decending paths in the spinal cord
               2) Spinal cord is also involved in reflexes and pattern generators
     2.  PNS
          a. Autonomic Nervous System (ANS, involuntary regulation)   
          b. Somatic Nervous System (involved in muscle contraction
              and voluntary behavior)
 
II.  AUTONOMIC: SYMPATHETIC & PARASYMPATHETIC

  A. Organization and general functions of this system
  B. Neuronal and functional organization of the ANS
     1. Organized as a two neuron system
          a. Differences exist for: 1) orgin of nerves, 2) postsynaptic
             neurotransmitter (NE verses Ach), and 3) position
             of ganglion
     2 . What does the ANS regulate? 
     3. The ANS is important in stress and other behavioral responses
          a. Response to an acute stressor: Cannon's "fight or flight" response 
          b. Response to a chronic stressor: CRH, ACTH, and Corticosterone
          
II. SOMATIC NERVOUS SYSTEM CONTROLS MUSCLE CONTRACTION **

  A. Somatic Nervous System
     1. Somatic nerves always innervate skeletal muscle
  B. Neuromuscular junction are only excitatory
     1. A unique mechanism occurs at the neuromuscular junction
          a. Ca++ initiates Ach release from presynaptic neuron
          b. Ach --> opens Na+/K+ channel (transmitter-gated) --> End Plate 
              Potential (EPP) --> opens Na+ channel (voltage-gated) -->
             Action Potential --> Muscle fiber contraction
  C. Motor unit
     1. Motor unit recruitment permits a graded response in  the muscle 
         organ
  D. Slow (Type I)  and fast (Type IIa and IIb) twitch muscle fibers
     1. Functional properties of Type I and Type II
          a. Fast twitch can be oxidative (IIa) or glycolytic (IIb)
          b. Significance of slow and fast twitch muscle fibers

IV.  MOLECULAR BASIS FOR MUSCLE CONTRACTION **

  A. Basic muscle microanatomy
     1. Muscle organ --> Muscle fiber (the cell) --> Myofibril --> Sarcomere
         --> Myofilaments (actin, myosin) are within the sarcomere
     2. Cytoarchitecture of a Sacromere--the basic contractile unit in 
         the myofibril
          a. There is functional organization of the myofibrils
              (actin and myosin)
  B. Molecular Basis for Muscle Contraction (Sliding filament mechanism)
     1. Myosin (thick filaments; site of ATPase activity)
     2. Actin (thin filaments)
     3. Tropomyosin and Troponin 
     4. T tubules and Sacroplasmic Reticulum (SR) -- 
         AP mediation
     5. Ca++ release from the SR initiates contraction
          a. Ca++ acts on Troponin to reveal cross-bridge 
              binding sites
          b.  Cross-bridge formation causes contraction of the muscle fiber
               1) ATP is utilized with cross-bridge formation
               2) Cross-bridge formation is repeated over and over (=cycling)
   C. Summary of these processes (Animation) 

V. REFLEXES AND CENTRAL PATTERN GENERATORS **

  A. What is a reflex and how are they important in behavior?
     1. Example 1: Withdrawal reflex
     2. Example 2: Crossed extensor reflex 
   B. Muscle Spindles and the Stretch Reflex
     1. Monosynaptic reflex
          a. Neural circuitry tends to maintain the status quo
     2. This circuit includes a:
          a. Primary afferent (senses stretch and rate of 
              change in stretch)
          b. Secondary afferent (stretch only)
          c. Alpha motor neuron (efferent which innervates
              extrafusal muscle)
          d. Gamma motor neuron (efferent which innervates
               intrafusal muscle)
     3. The CNS intervenes to elicit contraction and 
         motor behavior via alpha-gamma coactivation **
  C. Central Pattern Generators (CPG) and behavior
     1. The circuitry for some involuntary behaviors, 
         such as walking, reside in the spinal cord
          a. Early experimental evidence	   
          b. Role of the higher brain centers in control of 
              these spinal circuits
          c. Spinal cord circuitry responsible for lamprey swimming and  human walking 
              are similar
          d. Can the spinal cord learn?


VI. THE BRAIN CONTROLS MOST BEHAVIOR

  A. Voluntary behavior: Idea --> Program --> Execution --> Feedback
  B. Voluntary behavior involves many brain areas (Overview)
     1. Prefrontal cortex--Idea
          a. Retrieves relevant information from various cortices 
     2. Supplementary motor area (SMA)--Program
          a. Example of SMA function
     3. Premotor area (PMA)--Program
          a. Integrates numberous sensory inputs in guiding behavior
     4. Primary motor cortex (M1)--Execution
     5. Basal ganglia and Cerebellum--Feedback

  C. Motor output passes through one of several
      descending spinal tracts
     1. Some parallel processing exists
  D. Cortical input to the basal ganglia forms a loop back
       to the SMA, PMA, and M1
     1. Basal ganglia output is excitatory
  E. Cortical input to the cerebellum forms a loop back to M1
     1. The cerebellum monitors but does not execute behavior
          a. How does behavior change following a cerebellar lesion?
     2. The cerebellum has a number of functions
          a. Involved in balance and posture
          b. Has a memory for past behaviors
          c. Uses various sensory inputs to assess the status of a 
              executed behavior
          d. Modifies behavior via M1 **
  F. Summary on neural control of voluntary behavior

STUDY QUESTIONS

  1. What are the general functions of the autonomic and somatic nervous systems? How is the autonomic nervous system subdivided?

  2. Outline the neuronal characteristics of the sympathetic and parasympathetic nervous systems with regard to: 1) number of neurons involved, 2) location of the ganglion, and 3) anatomical origin of the system. See Figure 3.26 in Carlson.

  3. Starting with the presynaptic release of Ach, outline the sequence of cellular events which eventually results in an action potential on the muscle fiber.

  4. What is an end plate potential (EPP)? Compare and contrast with an excitatory post-synaptic potential (EPSP).

  5. Homeostasis of a number of physiological parameters, such as the rate digestion, is accomplished through opposition of the sympathetic and parasympathetic nervous systems. List an example where these two systems work in opposition. List an example of where they do not work in opposition. Is the parasympathetic nervous system always inhibitory? Is the sympathetic nervous system always excitatory? See Figure 3.26 in Carlson (Note: Sympathetic does not inhibit salivation as shown in this figure.)

  6. You are shown a nerve innervating an exocrine gland (e.g., a salivary gland). You decide that it is not part of the somatic nervous system. Why did you make that decision? What anatomical criteria might you use to determine if the nerve originates in parasympathetic or sympathetic nervous systems? What different pharmacological or physiological approaches (provide more than one possibility) might you employ to confirm your anatomical evidence?

  7. What is the difference between a muscle organ, muscle fiber, myofibril, sacromere, and myofilament (e.g., actin)?

  8. Outline the molecular mechanism for contraction of skeletal muscle. At what point is ATP used and for what purpose?

  9. What makes it possible to have a graded contraction of the muscle organ if each muscle fiber (=muscle cell) contracts in an "all or none" manner? Explain the inconsistency.

  10. The action potential spreading across the muscle fiber follows the "all or none" law. By what mechanism does the action potential elicit muscle contraction? Is the ratio within the sarcomere necessarily 1 action potential: 1 cross-bridge formation? Explain.

  11. Draw and label the cross-section of the spinal cord. Identify ascending neural tracts and descending neural tracts. What is the relationship between the spinal cord and the more peripheral spinal nerves? That is, where do peripheral nerves synapse in the cord? (See Figure 3.23)

  12. Draw a muscle spindle (include all afferent and efferent neurons). Which portion of the muscle is contractile? noncontractile? How does the stretch reflex work to maintain the status quo? How does outside motor input from the CNS modify the stretch reflex to permit muscle contraction?

  13. Why even have a stretch reflex?

  14. Read about the polysynaptic inhibitory reflex in the text. Under what circumstances is this reflex used?

  15. What is the minimum number of neurons involved in a simple reflex? Cite an example. What circuitry is involved in a simple reflex, such as the withdrawal reflex? Can the brain override some simple reflexes? Explain.

  16. You step on a thumb tack. Immediately, you withdraw your foot. First, trace the neural pathway inovolved in the immediate withdrawal. What compensation prevents you from falling? Second, in outline form, trace the ascending pathway involved in the realization that you stepped on something. What is the neural basis for localizing the stimulus (prick of the tack) to the foot and not to some other part of the body? Explain.

  17. It was once thought that the spinal cord was a structure that merely shuttled neural information between the periphery and the brain. Why is that idea now disputed? Explain.

  18. When you flex your forearm, motor input from the CNS results in contraction of the biceps muscle. But at the same time you extend the opposing triceps muscle which also contains stretch receptors that become stretched and thus should resist any lengthening. Propose a reflex mechanism that would explain this apparent contradiction. Draw a neural circuit that would explain these actions?

  19. What is the role of the cerebellum in movement? Play special attention to how the different cerebellar regions have specific functions in the control of movement. What is the role of the basal ganglia in movement? Compare and contrast cerebellum and the basal ganglia function.

  20. How are the followiing structures (lettered A and B) involved in mediating 1) motor output from the cerebral cortex and 2) motor input to the cerebral cortex? What are the functions of these structures in motor events?

  21. The electric organ of some electric fish consists of highly modified muscle cells, called electrocytes. Briefly compare the innervation of the electrocyte and the innervation of a muscle cell. How are they alike? How do individual electrocytes communicate with one another? For what purpose?

  22. Define and give the physiological significance (how or why it is important) for the following terms:
    • Motor unit
    • Troponin
    • Tropomyosin
    • Actin
    • Myosin
    • Sarcomere
    • Cross-bridge cycling
    • T-tubules
    • Ca++ (role in muscle contraction)
    • Sarcoplasmic reticulum
    • End plate potential
    • Putamen
    • Caudate nucleus
    • Striatum
    • Globus pallidus
    • Ventrolateral nucleus (of the thalamus)

  23. What are apraxias and what do they tell us about how the brain controls purposeful behavior? See Carlson.

  24. Read about Parkingson's disease and Huntington's disease in the text. What do these diseases tell us about how the basal ganglia function?

  25. Read about the major descending motor pathways (see Table 8.1 for a summary). Note that some functional overlap exists for these descending pathways. What might be the advantage of such parallel processing?

  26. What is a central pattern generator (CPG)? How are they important in control of animal behavior? What is the role of the brain in central pattern generator function? Give some examples of central pattern generators in humans and other vertebrates. How do these CPG's work?

  27. A voluntary behavior can be divided into the following components: Idea --> Program --> Execution --> Feedback. With this sequence in mind, explain the roles of the prefrontal cortex, premotor area (PMA), supplementary motor area (SMA), and primary motor cortex (M1) in carrying out a voluntary behavior. What are the roles of the basal ganglia and cerebellum in the execution of a voluntary behavior?

  28. After much practice a baseball pitcher learns to throw a good curve ball. What changes in brain function allow us to excel at complex behaviors following much repetition (especially ballistic behaviors which occur very rapidly, such as throwing a curve ball)? See the section on the cerebellum in Carlson.


ADDITIONAL INFORMATION ON THE INTERNET

Pharmacology and Physiology of the Autonomic Nervous System. A very detailed treatment of the subject from the Washington University School of Medicine in St. Louis. Links to related subjects.

Motor Systems Slide Presentation

Animations on muscle receptors and reflexes from the University of Western Ontario

Muscle Structure and Function in detail from the University of California at San Diego.

Network Patterns and Locomotion. A general article on pattern generators and vertebrate locomotion.



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