Motor System Spinal Reflexes Georgia Bishop, Ph.D. Professor and Vice Chair Department of Neuroscience bishop.9@osu.edu OBJECTIVES At the end of the module you will learn to: Describe the peripheral receptors and pathways that regulate spinal reflexes. 1. 2. 3. 4. 5. 6. 7. Define the terms proprioception and proprioceptor. Define the terms motor unit and recruitment. Explain how motor units function to increase muscle tension is increased Describe muscle spindles and Golgi tendon organs. Differentiate between alpha and gamma motor neurons. Describe the term Gamma Bias and explain its functional role. Differentiate the role of the muscle spindle and the Golgi tendon organ in proprioception. 8. Describe the neural correlates of spinal reflexes including the stretch reflex and flexor 9. withdrawal with crossed extension reflexes. 10. Describe the clinical significance of hyperactive or absent reflexes. 2 REFLEX CIRCUITS Reflex circuits in the spinal cord produce automated responses adaptive for typical situations. When a specific kind of sensory input consistently elicits a particular response, we call this a reflex. Spinal or reflex circuits govern many muscle recruitment patterns within and between limbs, including standing and walking. Reflex circuits require, at a minimum, 2 components: A sensory input A motor output In the circuit, the sensory neuron synapses on the motor neuron which then elicits a muscle contraction. Many reflex circuits also contain interneurons that may either excite or inhibit populations of motor neurons. Spinal Nerve Intervertebral Foramen Vertebral Canal Dorsal Root DRG Cell Dorsal Horn Periphery Dorsal Root Ganglion Dorsal Sensory Ventral Motor Spinal Nerve Ventral Horn Motor Neuron Ventral Root Efferent Axon of Motor Neuron Peripheral Process ALPHA MOTOR NEURONS 40 m Innervate extrafusal muscle fibers: Regular contractile portion of muscle SKELETAL MUSCLE MUSCLE FIBER Skeletal muscle is made up of bundles of cylindrical muscle fibers. Each muscle fiber is a single, multinucleated cell MOTOR UNIT Defined as a single alpha motor neuron and ALL THE MUSCLE FIBERS it innervates Each muscle fiber receives input from a single motorneuron which synapses at the single motor end plate. However, a motor axon may innervate more than one muscle fiber). MOTOR UNIT The size of motor units varies from small (10 – 100 fibers/motor neuron) to large (100 – several thousand fibers/motor neuron). Small motor units provide more precise control of motor activity. These would be found in muscles that control individual digits or muscles that control movements of the eye. MOTOR UNIT Motor End-Plate – Site where axons make synaptic contact with muscle fiber MUSCLE CONTRACTION MUSCLE TENSION 1 TWITCH (100 msec) The force of contraction of individual muscle fibers is determined by the firing frequency of the motor neuron Total force of contraction of a muscle is determined by number of alpha motor neurons that are active. MUSCLE CONTRACTION Tetany = a sustained muscular contraction caused by a series of stimuli repeated so rapidly that the individual muscular responses are fused. Maximal force a muscle can generate. Temporal summation. TETANIC CONTRACTION TETANUS Clostridium tetani is a grampositive rod-shaped bacterium that is found worldwide in soil; it is usually in its dormant form, spores, and becomes the rod-shaped bacterium when it multiplies. 1. 2. 3. 4. 5. Binds to peripheral nerve terminals and transported within the axon to CNS. Binds to proteins at presynaptic inhibitory motor nerve endings and taken up into the neurons. Effect is to block release of inhibitory neurotransmitters (GABA, glycine). Results in uncontrolled firing of motor neurons resulting in muscular spasms. Acts by selective cleavage of a protein required for neurotransmitter release, synaptobrevin II PROPRIOCEPTION AND PROPRIOCEPTORS PROPRIOCEPTION include awareness of the body’s position in space, sensation of forces acting on the body, and sensation of body movements underway Some define proprioception as position awareness and kinesthesia as awareness of movement. For this module, proprioception is used inclusively. A PROPRIOCEPTOR is a sensory receptor that is principally used for proprioception SENSORY INPUTS INFLUENCE RECRUITMENT Two main kinds of proprioceptors influence recruitment levels in the motor pools: Muscle spindles - muscle length, and velocity of muscle length changes Golgi Tendon Organs (GTOs) - muscle tension (force) information Cutaneous receptors, receptors in joints, and pain receptors also influence recruitment MUSCLE SPINDLES Muscle Spindle Muscle spindles detect muscle length, position, velocity, and acceleration Extrafusal fibers are the muscle fibers we have been talking about. They make up the muscles we see. Muscle spindles are miniature, long, thin stretch receptors that are present in all striated muscle. They are made up of specialized intrafusal muscle fibers surrounded by a capsule of connective tissue. They are scattered throughout the muscle and are aligned in parallel with the extrafusal fibers. SPINDLE EF The primary sensory afferent from the muscle spindle is called the Group Ia afferent (Ia) MUSCLE SPINDLES HAVE 3 MAIN COMPONENTS Intrafusal muscle fibers provide regulation of muscle spindle stiffness, for variable sensitivity to stretch Primary sensory axons (Ia and II) terminate on central region to sense stretch The type II afferents only detect length of a muscle whereas type Ia detect length and rate of change in length. Gamma motor axons synapse on intrafusal fibers to regulate their tension (stiffness) 16 TYPES OF MOTOR NEURONS Alpha motor neurons (medium – large neurons) Project to extrafusal (skeletal muscle) fibers Responsible for generating the muscle forces used to control movement Gamma motor neurons (very small neurons) Only present to control muscle spindles by synapsing on intrafusal muscle fibers Cannot produce any appreciable muscle force ALPHA-GAMMA CO-ACTIVATION (GAMMA BIAS) The CNS independently regulates gamma motor neurons GOLGI TENDON ORGANS Receptors found at the junction of the muscle fibers with the collagenous tendon. Unlike the muscle spindles that are arranged in parallel to the muscle fibers, the receptors in the GTO intertwine with the collagen fibers of the tendon. “Ib afferents” coming from the GTOs convey force data to the spinal cord. Detect muscle tension. 19 CIRCUITRY OF GOLGI TENDON ORGAN The circuit of the Ib afferent is: Activated Ib afferent synapses on inhibitory interneuron in the spinal cord. Ib excitatory interneuron This inhibitory interneuron inhibits the motor neuron that projects to the contracting muscle to decrease tension and prevent damage to the tendon. STRETCH REFLEX WITH RECIPROCAL INHIBITION As described previously, when a muscle is quickly stretched, Ia afferent synapses excite that muscle’s alpha motoneurons The is the stretch reflex Muscle length homeostasis The Ia afferent also synapses on the Ia inhibitory interneuron, which inhibits antagonist moto-neurons, relaxing that muscle. Provides a complementary functional effect. 21 STRETCH REFLEX Muscle spindles respond to weight of mug and activate the biceps to keep it upright. At same time, inhibitory interneuron blocks activity in the triceps. Increasing weight causes passive stretch of the muscle and increased activation of Ia afferent which activates more motor neurons to produce stronger contraction. May recruit more motor units. Increased recruitment of more motor units brings arm back to level position and overcomes increased resistance. FLEXOR WITHDRAWAL WITH CROSSED EXTENSION Results from activation of a nociceptor. Primary afferents branch into collaterals that activate 2nd order sensory neurons in the dorsal horn These then project to another set of interneurons in the spinal cord which have multiple effects. They project to: 23 1. Inhibitory interneurons that inhibit alpha motor neurons that project to the quadriceps (stance muscle) and to excitatory interneurons that activate flexor muscles (e.g., hamstrings). 2. Excitatory interneurons on the contralateral side of the spinal cord that activate the contralateral quadriceps and inhibitory interneurons that inhibit the contralateral flexor muscles. EFFECTS FROM JOINT RECEPTORS Joint receptors contribute some proprioceptive awareness, especially at end of the joint range and under high force or strain conditions Joint receptors can promote stability or safety Rapid reaction to high joint force is increased muscle tone around the joint Severe, acute pain from trauma causes extreme recruitment to splint the joint Long term pain and swelling produces inhibition of muscles to protect the joint CLINICAL APPLICATIONS Tendon jerk reflexes reveal CNS excitability state. When cerebral cortex is damaged, the default state of gamma motoneurons is hyperactive, and stretch reflexes are exaggerated. Absence of a reflex indicates there is damage at the level of the spinal cord. This may result from damage to a sensory neuron or the Ia afferent (e.g., diabetes) or damage to the motor neuron or pools of interneurons. Stretching exercises should be performed slowly less likely to elicit a stretch reflex. Smooth, coordinated movement requires good proprioception SUMMARY 1. Sensation and control of reflexes is required for normal, coordinated movements 2. Proprioceptors are sensory receptors dedicated to monitoring movements and forces. 3. Muscle spindles (intrafusal muscle) are miniature muscle fibers that are aligned in parallel with the extrafusal muscle and detect length and rate of movement of a muscle 4. Golgi Tendon Organs are proprioceptors that are entwined with the collagen of a tendon and detect tension in the muscle. 5. Alpha motor neuron innervate extrafusal muscle and gamma motor neurons innervate intrafusal muscle fibers. For proper function, both are co-activated. 6. Gamma motor neurons determine the sensitivity of the muscle to small changes in length. 7. Spinal cord reflexes may be elicited by proprioceptors or by cutaneous (nociceptors) receptors. 8. The stretch reflex elicits muscle contraction following stretching of a muscle. In addition to activation of the stretched muscle, the antagonist is inhibited via interneurons in the spinal cord. 9. The flexor withdrawal reflex with crossed extension involves pools of interneurons that induce removal of a limb from a painful stimulus and extension of the contralateral limb to maintain stability. 10. Absent reflexes may indicate spinal cord or motor neuron damage. Abnormal reflexes (e.g., hyperactive) are a sign of damage to higher parts of the nervous system. Thank you for completing this module Questions? Bishop.9@osu.edu Survey We would appreciate your feedback on this module. Click on the button below to complete a brief survey. Your responses and comments will be shared with the module’s author, the LSI EdTech team, and LSI curriculum leaders. We will use your feedback to improve future versions of the module. The survey is both optional and anonymous and should take less than 5 minutes to complete. Survey