Transcribed by Stephen Holt Neuroscience 33 Motor Reflexes by Dr.

Schiff

Feb 27, 2014

We went through a whole period of sensory physiology with me and with Dr. Speilman and Tuesday was sort of autonomic, possible involuntary motor pathways and today and tomorrow Im going to be talking about voluntary motor pathways. Pathways involved in deciding to do something and then doing it. But without even looking at the cortex, one of the things that I want to discuss is what happens down in the spinal cord; lets say you send a command to a level of the spinal cord that regulates a particular muscle and theres all sorts of things going on down at the spinal cord level, these are spinal reflexes that alter, modify this command. You dont just send a message to an alpha-motor neuron and say contract that muscle. Because if I send a message to, lets say I want to touch my nose. If I send the message to the biceps muscle to bend the elbow, how far am I bending the elbow? This far? This far? Or am I punching myself in the nose which would lead to blood all over the place and not a good thing. You might find it entertaining. So even when a command comes down the descending the motor pathway from the motor cortex, it gets modified. And what Im going to be talking about today are the reflex pathways at the spinal cord level that modify what the commands tell the muscles themselves. Now one of the things that go on here is that we have as you know in and around the muscle and joints we have proprioceptors: these are sensory transducers that get information about the current status or position of a joint or a part of the body. And what Im going to start with is the spindle. Remember muscle spindles consist of essentially 3 components. Heres a sort of a muscle spindle very schematically. This is a cell. Just as a skeletal muscle fiber is a cell this is a cell. And its a striated muscle at both ends. Sarcomeres, it contracts, the whole thing. The central area is afferent(sensory). And what youve got here is a sensor in this central portion of the spindle that measures how long this sensory portion of the spindle is. And it sends out afferent action potentials proportional in frequency to the length. Well Im oversimplifying slightly; its a little more complicated than that. Because there are 2 basic types of spindles and Im not going into their structure and architecture, but schematically in terms of function there are (in fact Ill draw another spindle here). And this is the sensory portion. The difference between these two spindles functionally is this. The 2 types of spindles are designated dynamic and static. The dynamic spindles, the afferent action potentials that are produced or the frequency of afferent action potentials that are produced depend on the rate of changethis is dynamic and this is static okin the static ones, well start with that because I said static and pointed to dynamic didnt I? In the static spindles the action potential frequency depends on the length of the sensory portion of spindle. So the action potential frequency depends on is determined by length. So if you have it this long, you get a certain amount of frequency of afferent action potentials, if you stretch this and hold it at a new longer length, somehow you get a higher frequency of action potentials and itll stay depending on the length.

Transcribed by Stephen Holt

Feb 27, 2014

The dynamic spindle is different. Here the action potential frequency is determined by the rate of change of the length, dt. In other words, while the length is changing, you get afferent action potentials produced. Once it reaches the new length and stays there, the action potential frequency goes down to zero. So this is only giving off sensory action potentials when the length is changing. And in fact only when its increasing. Whereas the static spindles give off action potentials all the time depending on the absolute length. Ok. Where are these spindles found? (drawing) Well basically theyre found in skeletal muscle. Heres a muscle going from here to there. And heres a muscle fiber, one of the cells of the muscle striated and all that. And heres another muscle fiber, ok. And where the spindle is located is attached to one of them off at an angle there is the central sensory portion and theres the other end. Now theres a bit more to it. Because within the spinal cord there are motor neurons. This portion of the cell here, the ends, the nonsensory portions of the spindles are striated muscle. And striated muscle doesnt do anything unless its stimulated by acetylcholine. Thats an old story. So in order for this to function as a striated muscle, you need a motor neuron, just as you do for the ordinary muscle fibers, you need motor fibers coming in. So heres a motor neuron, heres a motor neuron and the same for the others. Except these motor neurons are smaller than the motor neurons(smaller diameter axons, theyre myelinated just like the motor neurons that control this muscle fiber here). These of course are alpha motor neurons. If you recall Dr Sangines lecutres ages ago, (theyre on the midterm, you got to recall them!) the classification of myelinated fibers (alpha, beta, gamma, delta according to ranges of diameter), the alpha motor neurons are biggest; these are gamma motor neurons because theyre somewhat smaller. But functionally they act like motor neurons, they release Ach and so on and are myelinated. Now, how does this whole thing work? Well there are 2 ways of looking at this. One is, lets say this spindle here is attached from one muscle fiber to another. If that spindle is stimulated by action potentials on the gamma motor neurons, these contractile portions are going to contract(the striated muscle part of the cell.) But these are much weaker, much smaller cells than the main muscle fibers of the muscle. Just to toss in a few words for your vocabulary, these regular muscle fibers are referred to as extrafusal. And the spindles themselves are referred to as intrafusal. That goes back a couple of centuries and I have no idea why they are referred to that way but the terms stick. The extrafusal fibers are big and strong. These little bity spindly things, the intrafusal fibers, the cells with the spindles(the spindle cells) are not capable of budging these big guys significantly at all. So whats going to happen? What happens is, if you stimulate the gamma motor neuron, send some action potentials here, the striated muscle portion of these cells are going to contract. But this is attached to a big, strong extrafusal muscle fiber. This is attached to a big extrafusal muscle fiber. So this cant pull those at all; they wont budge. So if this contracts and this contracts what actually happens? Well, the sensory portion is going to be stretched. In other words, if the contractile portion of the spindle is stimulated, this contracts this way, this contracts this way with the ends of the fibers remaining

Transcribed by Stephen Holt

Feb 27, 2014

fixed. And so the central portion of the cell gets stretched. And what does the central portion of the cell do when it gets stretched? It sends afferent action potentials to the spinal cord. Either proportional to the new length or proportional to the rate of change of the length in the dynamic spindle. Now, where does the sensory fiber from this spindle go? Well, you assume the cell body is in the dorsal root ganglion, theres an axon going out to the spindle. To the sensory portion of the spindle. And then theres another branch of the axon going into the spinal cord into the dorsal horn and it synapses directly onto the alpha motor neuron of that same muscle. So what youve got here is the sensory portion of the spindle(draws); afferent fibers go into the spinal cord and synapse onto the alpha motor neuron. The alpha motor neuron goes to one of these extrafusal fibers. And stimulates it, theres a synapse. So heres an interesting control pathway. If you decide to contract the muscle, there are two ways you can do it, if you think about this. You can send the message down to the spinal cord to stimulate the alpha motor neuron directly and the alpha motor neuron will send an action potential along its axon to the muscle fiber; compulsory, mandatory transmission, you remember at that synapse? The muscle fiber contracts. Or you can do it this way. You can send the message down from your cortex to this area of the spinal cord and stimulate a gamma motor neuron going to a spindle within the muscle you want to contract. And what will that do? Well, it will cause the contractile portion of the spindle to contract, stretching the sensory portion of the spindle cell. The sensory portion of the spindle cell will send action potentials into the spinal cord which will synapse onto the alpha motor neuron, cause the alpha motor neuron to send action potentials to the extrafusal fiber and cause it to contract. This is a rather long way around. But it has an advantage. The advantage of your brain stimulating the alpha motor neuron directly is your muscle starts contracting fast, immediately. Because you send the command that goes directly to the alpha motor neuron, big axon, fast conduction, synapse to the extrafusal fiber, the muscle contracts. Its fast. The disadvantage is no message about when to stop. It just says contract. The alternative pathway is your cortex sends a message down a spinal cord to stimulate a gamma motor neuron, the gamma motor neuron causes the contractile portion of the spindle to contract which stretches the sensory of the spindle. The sensory portion of the spindle sends afferent action potentials that stimulate the alpha motor neuron and the alpha motor neuron causes the muscle to contract, but as the muscle contracts, this whole length of the spindle cell gets a little shorter. So what you did was you contracted the contractile portion of the spindle to stretch the center. But then, you shortened the length of the muscle a little bit which unstretched the sensory part of the spindle and then those sensory action potentials stopped and the alpha motor neuron is no longer being stimulated. So there is a limitation on how far that contraction goes, so I can now stop before I hit my nose. (student question) Ok, once the spindle contractile portion is stimulated, the contractile part of the spindle contracts this way. Which stretches the sensory part of the spindle. That sends action potentials that stimulate the alpha motor neuron.

Transcribed by Stephen Holt

Feb 27, 2014

The alpha motor neuron comes back to the extrafusal fibers and the muscle shortens. But this spindle the way it exists inside the muscle is a certain percentage of the muscle length. So if the muscle gets shorter, this overall spindle length from end to end gets shorter. So now youre unstretching the central sensory part of the spindle so itll stop sending out sensory action potentials and stop stimulating the alpha motor neuron so that the muscle will stop contracting. So this gives you a way of regulating how far the muscle is going to shorten when it contracts. Now, so in real life there are 2 ways of doing this, one is fast and one is more controlled. In real life, what do we do? And the answer is both. Theres a principle referred to as coactivation. Where when you decide, your primary motor cortex decides to contract your biceps muscle, it sends control message down the spinal cord to stimulate both the alpha and the gamma motor neuron. So, as soon as the alpha gets the message, the muscle starts to contract and its reinforced meanwhile by you stimulating the gamma motor neuron which stimulates the sensory portion of the spindle by stretching it. And that further stimulates the alpha motor neuron. So youve already had the advantage of speed, but part of this activation of the alpha motor neuron depends on the sensory input from the spindle so as the muscle shortens, that will go away. So youre still getting a certain amount of stimulation directly to the alpha motor neuron but itll stop being reinforced so youre actual contraction gets weaker and weaker and weaker and at some point itll stop. So the whole point here is that you by coactivation you have the advantage of control as well as the advantage of speed. But this pathway, this kind of back and forth pathway, leads to an interesting situation. Suppose you have a way of mechanically stretching a muscle, suppose you stretch your muscle, you grab onto the tendon and pull. Well that requires surgery. But remember the quadriceps muscle anterior to the thigh. The quadriceps tendon loops over the patella and inserts on the tibia. So as that muscle contracts the knee straightens because its pulling that tendon. But suppose, what you do is take the little triangular headed rubber mallets and you go bang onto the patellar tendon. Now what does that do? Well it stretches the quadriceps muscle slightly. Whats that going to do? Its going to stretch the spindles in that muscle. Its not a major stretch so the static spindles wont be affected by too much, but the dynamic spindles will suddenly see a change in length, theyll start discharging action potentials which synapse onto the motor neuron(the alpha motor neuron) and cause this muscle to contract. So you hit the patellar tendon with a hammer and you get a little kick. And thats a good way to check whether a persons pathways are intact because there are a number of conditions, by way of nerve damage or other conditions where you dont get a kick or get a weak kick. Or under certain specific circumstances you might get a pendullar kick, where it starts swinging and just keeps swingingwell get to that tomorrow. So the whole idea is when you stretch a muscle, you can do it without that sort of thing. If I hold my hand out and Im not really looking and you drop a small kilogram weight onto my hand, itll initially stretch the biceps muscle which will cause a reflex contraction and Ill end up keeping my hand more or less steady.

Transcribed by Stephen Holt

Feb 27, 2014

Again, when this happens, its referred to as a myotatic reflex. And the most interesting thing about it, and why its worth discussing at all is that how many synapses are involved? Just the one, well 2 if you count the neuromuscular synapses from the alpha motor neuron to the extrafusal fiber. But you stimulate the spindle as one synapse and then you send action potentials to cause the muscle to contract. And because any pathway that has multiple synapses in it takes longer because there is amount of delay of transmitter release, transmitter has to diffuse across the synaptic space and bind to receptors and then if the second receptor is metabotrophic that starts the whole thing of activating G proteins and such, time is lost. Possibly a quarter to a half millisecond. And so what happens here is this is the fastest reflex around because theres only one synapse; you cant get less than one synapse(plus the neuromuscular synapse). But the key here is one other thing, and that is whenever a muscle length changes, the muscle is stretched as in the patellar tendon being hit by a hammer, theres a feedback loop initiated, the length goes up, that stimulates the alpha motor neuron cause it stretches the spindle, afferent fibers stimulate the alpha motor neuron, the alpha motor neuron causes the extrafusal fibers to contract and the length comes back to itsthe muscle shortens again back to where it had been originally. So what this whole reflex pathway here does is it provides negative feedback; regulation of muscle length, thats important. Because it enables you to sense the length of the muscle at some level and keep your hand or whatever limb in the same position. Cause if anything changes theres this feedback loop from the spindle that will restore the original position. Second important pathway thats involved in feedback regulation of muscles at all is this. (draws) And this involves a structure known as the golgi tendon organ named for the guy who first got a picture of it. And where is the golgi tendon organ found? In a tendon, duh. So heres your muscle, here its anchored at the origin, the insertion is by way of a tendon, so theres a tendon. And embedded in the tendon is this structure which is the golgi tendon organ. What does it do? Its a stretch receptor effectively. The cellular structure, this transducer, is actually a stretch receptor, but whats it measuring the stretch of? Measuring the stretch of the tendon its embedded in. So if the tendon gets stretched, the golgi tendon organ gets stretched and it sends afferent action potentials out signaling that its been stretched. But how do you stretch a tendon? You have to apply force. And a tendon is essentially an elastic, its not an active contracting organ, rather the tendon is a spring, a rubber band, an elastic piece of material. Im sure Dr Craig has described all the wonderful parts that go into its structure. And you dutifully memorized them and thank god I dont have to. But the whole point is that when you have an elastic piece of tissue like this tendon and you have this golgi tendon organ embedded in it, the amount of stretch(change in length) is proportional to the amount of force thats exerted on it. Because in order to stretch it you have to apply force. And this is a passive spring, or elastic. So it has to follow Hookes Law where the change in length is proportional to the change in force or tension. See, thats probably the oldest guy Ill mention by name, were talking 17th century. So the sensory discharge measures how much the tendon has been stretched and therefore indirectly measures how much force there is on the tendon, ok. Now, what affects

Transcribed by Stephen Holt

Feb 27, 2014

the force on the tendon? Well, when the muscle contracts it pulls on the tendon. At the same time, if the muscle stays at a certain degree of contraction and then somebody pulls on the joint, forcibly extending the biceps, that will stretch the tendon also. And so it doesnt matter if its active force or active tension or passively applied by someone else pulling your leg. But, the golgi tendon organ sends afferent discharges proportional to tension in the muscle. And the other factor here is that the afferent fibers from the golgi tendon organ probably by way of a little interneuron(so theyre a little slower) inhibits the alpha motor neuron. What does that mean? The alpha motor neuron determines how much force the muscle is contracting by sending out whatever frequency of action potentials that will cause the muscle to contract. That generates tension which stretches the golgi tendon organ which signals there is a certain amount of tension and inhibits the alpha motor neuron. So this is a negative feedback loop regulating muscle tension. So, the golgi tendon organ is negative feedback regulation of tension. Now theres one situation in which these things sort of come together, and thats what known generally as the clasp knife reflex. And that is if youve got someone who is unconscious or recently dead or in other words not paying attention to a particular joint, lets say your elbow. And you passively bend this persons elbow, what are you doing? Well theres an extensor muscle here, the triceps, which you are elongating as you bend it. So the spindles in the triceps will send afferent fibers, afferent action potentials, to the alpha motor neuron of the extensor muscle and cause the extensor muscle to contract, thats your myotatic reflex. And so the extensor muscle will resist your attempting to bend the elbow and you keep pushing and that does more and more; stretching the extensor muscle, the extensor muscle contacts to try not to be stretched. But at the same time the amount of tension in the tendon of the extensor muscle is gradually increasing because youre applying force and the muscle is contracting more. And you apply more force, elongate it, and the muscle is applying more force. And so the force exerted by the extensor muscle in this case is gradually going up as a result of the myotatic reflex because youre stretching the muscle and that causes the spindles to signal that its being elongated and therefore it stimulates the alpha motor which tries to straighten it. So theres more and more tension in the muscle and this is your myotatic reflex triggered tension going up. Now, the discharge, afferent discharge of the golgi tendon organ down here is not very large but eventually you start really stretching that tendon and putting tension on that tendon and the golgi tendon organ is going to start discharging and remember its inhibiting the alpha motor neuron. So the golgi tendon organ discharge, this is spindle discharge will at some point exceed the stimulation of the muscle by the myotatic reflex. And what then happens is it resists, resists, and then suddenly you reach a point where suddenly the golgi tendon organ inhibits the alpha motor neuron more than the spindle is stimulating it and the muscle relaxes and stops resisting and this just goes. Its called the clasp knife. Clasp knife is one of the terms used for a pocket knife basically. Have you ever closed a pocket knife that folds? You fold the blade and it resists for a while

Transcribed by Stephen Holt

Feb 27, 2014

and then it just snaps close. And hopefully the bow of your thumb of your other hand isnt in the way. Its called a clasp knife reflex. Thats basically an accommodation of these first two reflexes. Theres one more thats important. And thats whats known as the Renshaw cell pathway. Heres your half a spinal cord. We used to have people in the anatomy dept who draw a whole spinal cord with 2 hands, but nah, they all go to power points and plastinations. Heres the ventral cord of the spinal cord and youve got an alpha motor neuron here and its sending an axon out to the muscle. This axon branches and sends a little branch out entirely still within the gray matter of the spinal cord. And this synapses onto a little neuron thats present entirely in here(Ill put a letter R in here for Renshaw cell). And the Renshaw cell sends a little axon back and inhibits the alpha motor neuron. This is excitatory synapse plus it inhibits the alpha motor neuron. So think about this: if various other pathways stimulate the alpha motor neuron, so the alpha motor neuron sends out an action potential, that action potential is already on its way to the muscle. At the same time, it stimulates the Renshaw cell which inhibits the alpha motor neuron so that alpha motor neuron becomes less likely to produce a second action potential. Why would you care about that sort of thing? Well the reason is this. Every muscle in your body is made up of what is called motor units. A motor unit is all the muscle fibers that are innervated by and controlled by the same alpha motor neuron because alpha motor neurons when they get near the muscle start to branch and so one alpha motor neuron might regulate control 2 or 3 or 6 or 100 muscle fibers. And obviously since transmission from the alpha motor neuron to the muscle fiber either happens or doesnt, its mandatory transmission, if you stimulate an alpha motor neuron youre going to stimulate that 10 or 100 muscle fibers. Ok. Youve got a whole bunch of alpha motor neurons in this ventral horn that each stimulates its own motor unit. Its own subset of one particular muscle. So if you want to contract that muscle, you send a command down that will excite/depolarize all of those alpha motor neurons. Well which of those alpha motor neurons will initiate action potentials and cause their muscle fibers to contract? Well, lets just say its whichever ones have the lowest threshold and are the easiest to excite. So some are more easy to trigger and some not so easy. So the bottom line is that if you didnt have any further regulation going on, the alpha motor neurons and their muscle fibers that have the lowest threshold would do all the work and the ones with a slightly higher triggering threshold would never get excited. Now, thats not very practical because youre going to have some of the fibers in a particular muscle that get exhausted and the rest of them are just on vacation. So what this Renshaw cell reflex does is this: you excite(this is the lowest threshold alpha motor neuron). So you trigger an action potential in it, it excites the muscle fibers in its motor unit. At the same time it stimulates a Renshaw cell which inhibits that alpha motor neuron so thats now a little hyperpolarized(has an IPSP in it.) And then what? Well the next time your brain sends a stimulatory message down to that muscle, thats no longer the easiest alpha motor neuron to excite because that alpha motor neuron has been hyperpolarized, has an IPSP around it and they last a long

Transcribed by Stephen Holt

Feb 27, 2014

time. So the next stimulatory attempt thatll have a higher threshold, itll be harder to stimulate that alpha motor neuron then a different one, but of course it has the same pathway with its own Renshaw cell so it will stimulate its muscle fibers and at the same time trigger a Renshaw cell to inhibit it so it will be hyperpolarized and harder to excite. So essentially what it does is it forces the alpha motor neuron and the different motor units within a muscle to share the work. In the years PP(preplastinations) when you actually learned anatomy on cadavers, there would be 6 or 8 students around one horizontal person and it would always be one that did all the dissecting while the others stood around and watched. And that was the analogy I always gave. It forced that person after he made a cut to go to the back of the line and let someone else do the dissecting. Ok. So these are the major reflex pathways involved just in stimulating an alpha motor neuron and its motor unit to contract. Theres feedback regulation of length, theres feedback regulation of tension. Usual example of that is you can pick up an egg without crushing it. But there are also other reflex pathways that involve more than just half of a spinal cord or half of a ventral horn of a spinal cord because there are reflexes that involve both sides of the body. For example, if something stimulates the flexor in one of your legs, theres going to be a reflex activation of the extensor muscle in your other legs so you dont fall down. And this goes back in forth from side to side when you walk. So what we end up doing is when you repeatedly do something that involves alternating the sides of your body, what gets set up in the spinal cord is whats known as a pattern generator. The reflexes from one leg stimulate the opposing muscles in the other leg and they alternate back and forth, back and forth, setting up a pattern of, for example, walking. Lets go back to something you had when you were looking at the spinal cord, the Brown-Sequard syndrome. Remember when you have half of a cut in a spinal cord confined to one side, the amazing thing about these people is that they can walk. Not over hilly ground perhaps and not on a path that requires a lot of turns, I mean crossing 1st ave at 23rd St is a problem. But I mean as long as theyre walking on level ground in a straight line they can walk. Because what happens is the pattern generator for walking is established in the lumbar cord and all you have to do is send down a signal in whichever side of the spinal cord is still intact in the Brown-Sequard person that says start. And then the walking reflexes go automatically as a result of the pattern generator. But, why dont we take a 5 minute break now and Ill move on to the descending motor pathways and the like.