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CHAPTER _13.docx

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Brock University
John Mitterer

INTRODUCTION The motor system consists of all our muscles and the neurons that control them. The spinal cord contains certain motor programs for the generation of coordinated movements and that these programs are accessed, executed, and modified by descending commands from the brain. Motor control is divided into two parts:  The spinal cord’s command and control of coordinated muscle contraction  The brain’s command and control of the motor program in the spinal cord THE SOMATIC MOTOR SYSTEM The muscles in the body can be described according to two broad categories:  Smooth o Lines the digestive tract, arteries and related structures o Innervated by nerve fibers from the ANS o Plays a role in peristalsis and the control of blood pressure and blood flow  Striated – there are two types of striated muscles o Cardiac muscle  is the heart muscle  contracts rhythmically  innervations of the heart from the ANS functions to accelerate or slow down the heart rate o Skeletal muscle  Constitutes the bulk of muscle mass in the body  Functions to move bones around joints  Is enclosed in a connective tissue sheath that forms tendons  Contains hundreds of muscle fibers, each innervated by a single axon branch from the CNS (see Fig. 13.3)  Derived from 33 paired somites  Controlled by the somatic motor system  Under voluntary control and it generates behaviour Consider elbow joint (see Fig. 13.2)  Movement in the direction that closes the knife is called flexion o The muscles that cause flexion are called flexors. Because they work together, they are called synergists.  Movement in the direction that opens the knife is called extension o The muscles that cause extension are called extensors.  Flexors and extensors pull on a joint in opposite directions and are antagonists. Muscles that are responsible for movements:  of the trunk are called axial muscles o important for maintaining posture  of the shoulder, elbow, pelvis, and knee are called the proximal muscles o important for locomotion  of the hands, feet, and digits are called the distal muscles o important for specialized manipulation of objects THE LOWER MOTOR NEURON Somatic musculature is innervated by the somatic motor neurons in the ventral horn of the spinal cord (see Fig. 13.3).  These cells are sometimes called lower motor neurons to distinguish them from the upper motor neurons of the brain that supply input to the spinal cord.  Only the lower motor neurons directly command muscle contraction. The Segmental Organization of Lower Motor Neurons The axons of lower motor neurons bundle together to form ventral roots. Each ventral root joins with a dorsal root to form a spinal nerve that exits the cord through the notches between the vertebrae. Skeletal muscles are not distributed evenly throughout the body, nor are lower motor neurons distributed evenly within the spinal cord. For example, innervations of the more than 50 muscles of the arm originate entirely from spinal segments C3-T1. The ventral horns in this region of the spinal cord appear swollen to accommodate the large number of motor neurons that control the arm musculature (see Fig. 13.4). The lower motor neurons are also distributed within the ventral horn at each spinal segment in a predictable way, depending on their function (see Fig. 13.5). Alpha Motor Neurons There are two categories of lower motor neurons of the spinal cord: alpha and gamma motor neurons. Alpha motor neurons trigger the generation of force by muscles.  One alpha motor neuron and all the muscle fibers it innervates collectively make up the motor unit (the elementary component of control).  Muscle contraction results from the individual and combined actions of these motor units. The collection of alpha motor neurons that innervates a single muscle is called a motor neuron pool. Graded Control of Muscle Contraction by Alpha Motor Neurons The nervous system uses several mechanisms to control the force of muscle contraction in a finely graded fashion.  The first way is by varying the firing rate of motor neurons. o An alpha motor neuron communicates with a muscle by releasing Ach at the neuromuscular junction. o Ach released in response to one presynaptic action potential causes an EPSP in the muscle fiber large enough to trigger one postsynaptic action potential  This will cause a twitch in the muscle o Sustained contraction requires a continual barrage of action potentials. High frequency presynaptic activity causes temporal summation of the postsynaptic responses. Twitch summation increases the tension in the muscle fibers and smoothes the contraction (see Fig. 13.7).  The second way is by recruiting additional synergistic motor units. o The extra tension provided by the recruitment of an active motor unit depends on how many muscle fibers are in that unit.  In the antigravity muscles of the leg, each motor unit tends to be quite large, with an innervations ratio of more than 1000 muscle fibers per single alpha motor neuron  In contrast, the smaller muscles that control the movement of the fingers are characterized by much smaller innervation ratios.  In general, muscles with a large number of small motor units can be more finely controlled by the CNS o Most muscles have a range of motor unit sizes, and these motor units are recruited in the order of smallest first, largest last.  This explains why finer control is possible when muscles are under light loads than when they are under greater loads. Inputs to Alpha Motor Neurons Alpha motor neurons excite skeletal muscles. Lower motor neurons are controlled by synaptic inputs in the ventral horn. There are three major sources of input to an alpha motor neuron as shown in Fig. 13.8. Types of Motor Units The red (dark) muscle fibers are:  Characterized by a large number of mitochondria and enzymes specialized for oxidative energy metabolism.  Slow to contract but can sustain contraction for a long time without fatigue  Typically found in the antigravity muscles of the leg and flight muscles of birds that fly The pale (white) muscle fibers:  Contain fewer mitochondria  Rely on anaerobic metabolism  Contract rapidly and powerfully, and also fatigue rapidly  Involved in escape reflexes Each motor unit contains muscle fibers of only a single type.  Fast motor units – contain rapidly fatiguing white fibers o The motor neurons are generally bigger and have larger-diameter, faster- conducting axons o Tend to generate occasional high-frequency bursts of action potentials  Slow motor units – contain slowly fatiguing red fibers o The motor neurons have smaller-diameter, more slowly conducting axons o Have relatively steady, low-frequency activity Neuromuscular Matchmaking Which came first, the muscle fiber or the motor neuron? This question was addressed in an experiment by John Eccles in which the normal innervation of a fast muscle was removed and replaced with a nerve that normally innervated a slow muscle (see Fig. 13.9).  It resulted in the muscle’s acquiring slow properties (i.e. type of contraction and its biochemistry). This change is referred to as a switch of muscle phenotype. Besides the alterations imposed by pattern of motor neuron activity, muscle fibers are also changed simply by varying the absolute amount of activity.  A long-term consequence of increased activity is hypertrophy of the muscle fibers  A prolonged inactivity leads to atrophy of muscle fibers EXCITATION-CONTRACTION COUPLING Muscle contraction is initiated by the release of ACh from the axon terminals of alpha motor neurons. ACh produces a large EPSP in the postsynaptic membrane due to the activation of ACh receptors which is sufficient to evoke an action potential in the muscle fiber.  By the process of excitati2+-contraction coupling, this action potential (excitation) triggers the release of Ca from an organelle inside the muscle fiber, which leads to contraction of the fiber. 2+  Relaxation occurs when the Ca levels are lowered by reuptake into the organelle. Muscle Fiber Structure – See Fig. 13.10  Fusion of myoblasts in early development leads to the formation of multinucleated muscle fibers.  Muscle fibers are enclosed by an excitable cell membrane called the sarcolemma.  Within the muscle fiber are myofibrils which contract in response to an action potential sweeping down the sarcolemma. 2+ o Myofibrils are surrounded by the sarcoplasmic reticulum (SR) (stores Ca ). o Action potentials sweeping along the sarcolemma gain access to the SR by way of tunnels called T tubules. o When the T tubule comes in close apposition to the SR, there is a specialized coupling of proteins in the two membranes. o A voltage-sensitive cluster of four calcium channels (tetrad) in the T tubule membrane is linked to a calcium release channel in the SR (see Fig. 13.11). o The arrival of an action potential in the T tubule membrane causes a conformational change in the tetrad, which opens the calcium release channel in the SR membrane.  This leads to an increase in free Ca2+within the cytosol which causes the myofibrils to contract The Molecular Basis of Muscle Contraction – See Fig. 13.12 The myofibril is divided into segments called sarcomeres by disks called Z lines.  On each side of the Z lines is a series of bristles called thin filaments. Thin filaments form adjacent Z lines face one another but do not come in contact.  Between the two sets of thin filaments are thick filaments Muscle contraction occurs when the thin filaments slide along the thick filaments, bringing adjacent Z lines toward one another. Thus, the sarcomere becomes shorter in length. This adheres to the sliding-filament model (see Fig. 13.13). The sliding of filaments occurs because of the interaction between the major thick filament protein (myosin) and the major thin filament (actin). The exposed “heads” of the myosin bind to actin and then undergo a conformational change that causes them to pivot (see Fig. 13.14). Now, the thick filaments move with respect to the thin filament.  ATP is required to disengage the myosin h
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