Transcribed by _Jessica Li Neuroscience Lecture 25 Vestibular System by Dr.

Schiff

February 16, 2014

Dr. Schiff Ok so now were going to go over the other job that hair cells have. And that is your vestibular organs and vestibular system. And what you have, what they are associated with, anatomically associated with the ear, the cochlea. And in fact the same endolymph that circulates through the cochlea is going to have an effect on the vestibular system as well. So um , what the vestibular system does is, is it helps keep you oriented in space. So that if youre turning, if youre accelerating, if youre in a car thats suddenly stops or starts or accelerates you can tell, detect what these changes in motion are. And there are two main components to the vestibular system. One is the labyrinth. One is the utricle. To be sure, the utricle is a flat organ that generally if you tilt your head slightly forward, horizontal. Theres another organ that is at right angles to it, smaller, thats also called the saccule. Were not exactly sure what the saccule does in human. In cats, it seems to be involved in balance and if you tossed a cat off the table on the floor it would land on its feet. They can do these maneuvers to become right side up. But for humans, its not clear what the saccule does. Some people have suggested that it detects vibrations but its not the best design, or the best arrangement to do that. And it doesnt do it very well. So well just ignore the saccule for the moment. And well go with the utricle and the labyrinth. The labyrinth consists of three semicircular canals. Which again is mislabeling at best because they are not semicircles they are full circles. And Im going to be talking about the labyrinth and the semicircular canals first and the utricle afterwards. What the canals do in the labyrinth is detect when you are turning; if youre turning, if youre rotating your head. This is very useful among our ancestors. Our cousin the orangutan swings from tree to tree. Lets go here and go through various maneuvers in space to grab the next branch of the next tree. And part of it is when your body moves you have to keep your eye aimed at the next branch you reach to. So one of the things that the vestibular system, particularly the labyrinth, helps you do is keep your eyes fixated on whatever target youre looking at. Now a more modern version of the orangutan story swinging through the trees is you riding down a road with all these potholes and youre trying to read a street sign about half a block ahead. And your body and head are being thrown left and right, up and down constantly, but you are able to keep your focus on the street sign and manage to read it. So if thats a better goal than getting through the forests of Borneo, then I dont know.

Transcribed by _Jessica Li

February 16, 2014

The semicircular canals have an anterior circular one, on each side of your head. Anterior, a posterior vertical canal, anterior vertical canal thats towards the front of your head and towards the back of your head. And theres also a horizontal semicircular canal. So among them they set up a sort of a coordinate system and can detect rotations of your head in any direction. But if you look at them anatomically, and its very hard to draw these fellows in 3D in a 2D blackboard, what youll see is that the vertical canals have a sort of common part piece of tubing. So you have 1 canal going this way and one canal sorta this way, and the horizontal canal is sorta in and out of the plane of the blackboard like that. So the key is, 3 canals, the 2 vertical ones have a length of tubing, plumbing, in common. So that if you were to do something to set, and they are filled with endolymph, and if you were to do something that would set the endolymph here moving, then that would start the endolymph here moving because they are passing though a common duct. So the anterior and posterior vertical canals cant be functionally separately completely. And for experimental purposes, clinical purposes, clinical diagnostic purposes, you generally tend to focus on the horizontal canal because that is not interfered with by any of the others. Now what you have here, if we just look at the horizontal semicircular canal, and as I said theyre not semicircles they are circles, is, Ill try to draw this sort of in the plane of the blackboard. This looks narrower because its further away and it gets a little lighter because were getting closer to the foreground. So picture this as going into the blackboard and theres an enlargement in the canal called the ampula. And this canal is surrounded by bone and lined by endothelium and filled with endolymph, the same endolymph that you had in your cochlear duct. High potassium solution and bathes the cilia, the hair cells, which Ill get to in a moment. The enlargement or the ampula, which I will enlarge here. For the most part, the whole system is lined by tissues, by cells. But here you have a projection of the cellular structure going into the middle of the lumen of the ampula. And this structure here, this projection forward is called the crista. C-R-I-S-T-A. And the crista has in addition to the structural cells that make up the wall of the lining of the semicircular canals has two additional types of cells. One of which is hair cells. So you have hair cells with their cilia projecting up. And all the hair cells in the crista of one semicircular are orientated the same way. They all have their tallest cilium to the left or the tallest cilium to the right. In other words, anything that moves them to the left or right will depolarize or hyperpolarize them all. Sp heres your semicircular canal and your loop completes into the blackboard. And the other kind of cell present in the crista are secretory cells that produces sort of Mucin or mucous or gel that projections into the lumen of the semicircular canals. Sp that you have this gel, this plug in the semicircular canals. This is called the copula. The copula ensures that the cilia of all the hair cells are either pushed left or 2

Transcribed by _Jessica Li

February 16, 2014

right in unison. So if the copula is bent this way all the cilia are bent that way. It keeps them synchronized. The other thing that is characteristic of the vestibular system hair cells is this: these hair cells are producing afferent action potentials in their sensory neurons when there is no stimulation. Theres no movement of the cilia but theyre still releasing some transmitter and creating some EPSP theyre generating action potentials at a very slow rate at about 5-10 ms. In here is endolymph. So what happens? Lets look at a glass of water. And just to make sure you can see the water were going to take a pepper mill and sprinkle some pepper flakes on the water so you can it. Water is standing still. Glass is standing still. Suppose I take this glass and turn it. What does the water do? The glass turned clockwise and the water stayed still. From the point of view of the glass the water moved counterclockwise. Lets take a look at the semicircular canals: the crista, the copula, the hair cells, cilia, etc. So lets say you turn your head a little bit. Generally the endolymph stays still just like my glass, the water stayed still. The ampula stays still. Youve go the endolymph stays still and the canal turns. The endolymph lags behind its going to push on the copula and bend the cilia. There are two possibilities: your brain cells this copula got pressed this way (to the left). What does this mean? The whole canal got moved clockwise and the endolymph stayed still or the head is perfectly still and the endolymph started moving. And moved clockwise and pressed against the copula there. There are two possibilities: which is true? The function of the vestibular system is to correct small movements of the head. Your eye moves in the opposite direction and keeps the target in focus. Its very rare to get movement of endolymph. Whenever your vestibular system detects a relative movement of endolymph pushing on the copula, and bends the cilia, that you generally dont make big moves and can assume safely that the endolymph stays still. Just like the water in the glass the endolymph stays still. What does this do? One thing it does is this: you have information from the vestibular system, feeds to the 8th nerve nuclei, they talk to each other. They set a, they send ascending messages to detect if anything is going on they send messages to nuclei to cranial nerve 6. Which has a subnucleus in it that coordinates all of this sort of stuff. And 6 then sends signals to 4 and also on to 3. What do these have in common (6, 4, 3)? They move the extraocular muscles in your head. And the whole idea is that is that the vestibular system detects that you have moved your head it signals the cranial nuclei 6, 4, 3 to move your eyeballs in the socket the other way to keep you looking at whatever you were looking at before.

Transcribed by _Jessica Li

February 16, 2014

So your eye moves compensates for any head movements. If I were to turn my head to the right my eyes automatically turn to the left to keep the same target fixated. Brain is making the assumption that the endolymph isnt moving. Your brain is usually right. Youll see some errors in a minute. If I slowly keep turning, people in general dont spin. Most people dont spin or the endolymph isnt moving. I turn to the right and my eyes move to the left. Unlike a dummy, my eyes cant keep turning to the left to make a complete circle. What happens is that there is a limit to how far the eyes can move to the left. As I continue to move to the right, the eyes will make the jump to a new focus point. It will turn left, left, left, left, and then switch to the right. Thats because your vestibular system detects the movement and tries to correct it by the slow leftward movement as Im turning to the right but cant do it so sometimes needs to do the fast jump to the right. When that happens its call nystagmus. In the specific example I have, by convention, causes right nystagmus. The nystagmus is named for the direction of the fast jump. If you think about it you have 3, 4 pairs of cranial nerve nuclei that make your eyes turn to the left as you turn to the right. So thats the major project. The fast jump to the right is a phenomenon that happens because theres a limit to how much this slow compensatory mechanism can handle. Its named for, by convention, the direction of the fast jump. But when your eyes look to the left and jump to the right is called right nystagmus. So lets see what happens when you slowly try to turn to the right. You eyes move slowly left and jump to the right, right nystagmus. If you were to start turning to the left slowly, you would develop left nystagmus because thats the fast jump. What happens if you keep turning? Just like this glass if you make little movements the water stays still. If you start this glass on the turntable, and keep it going, eventually the water, this friction between the glass and the water, its gonna catch up. Eventually you reach a point where if you have a glass moving at a steady rate the water within the glass is moving at a steady rate. Compare that to the horizontal semicircular canal. If you keep rotating, rotating, and rotating, eventually the endolymph will be moving along with your head. Whats the friction produced between this water and this glass? Vs how much water do you have in a glass. Here you have a donut shape too. More surface area per volume. A lot more friction so it takes very little turn (maybe 1 full turn) to get the endolymph moving. How does the nervous system react? Your turning but the endolypmh is moving with you so in what direction is the copula? Straight up. What are the sensory messages are you getting from your vestibular system that youre not rotating. And are you producing any kind of nystagmus? No. because your eyes are just moving along steadily. Theres no nystagmus.

Transcribed by _Jessica Li

February 16, 2014

Now lets do this for a while and then stop. You turn a turn or two and stop. At this point your endolymph is still moving. The endolymph sends a signal that either the endolyphn is moving clockwise to the right or your head is moving to the left. Your brain has learned that since you were a wee tod. Your brain concludes that at this point youve already been rotating to the left and stop, concludes that youre rotating to the left though you are not. Produces a left nystagmus. This is called a post rotational nystagmus. After the rotation has stopped. Post rotational nystagmus is always in the opposite direction of what the rotation has been. It doesnt take a lot of spinning. One turn can do it. Cant see this in a mirror; can only see on another person. Because your eyes can only see your eyes in the mirror so while theyre moving they cant see anything else. So post rotational nystagmus. If you move to the right you get right nystagmus. If you stop turning to the right you get left nystagmus. Which is post rotational form. The other thing that it does is that there are descending pathways, referred to as vestibulospinal tract which adjust your rostral muscles, your leg extensors and so on. So if your vestibular system detects that you are turning it may want to increase the tone of your extensor muscles in your other leg (in the lef thats going forward). The vestibulospinal tract is bilateral. You have cells that cross here and go to the other side of the spinal cord. One thing you can do to test the function of the semicircular canals is stimulate in some way movement of the endolymph and see if that induces the appropriate nystagmus. Because that will indicate if that whole ascending pathway is functioning correctly. How would you try to get endolymph rotating in one or both semicircular canals? One way to do it is thermally. This is referred to as caloric stimulation. Lets say you have a subject laying on his back, facing up, because hes lying down his horizontal semicircular canals are vertical. Then lets say you put water at room temperature into his right ear canal? Endolymph here is warmer that endolymph here because you warmed it up. What happens? You start conduction current. Like the way air circulates in your home when theres a cold window and you have a hot radiator and you get the heat rising. This endolymph goes up and this endolymph goes down and you get a circulation, counterclockwise to the endolymph in that canal caused by the warm water. What does the person consciously or unconsciously feel? If you assume that the endolymph is standing still then my head is turning clockwise. So he feels as though he is turning this way. And what will that produce? Right nystagmus.

Transcribed by _Jessica Li

February 16, 2014

So warm water, face up, right ear produces right nystagmus. With a little bit of thought you can see that it leads to 8 possible exam questions because this is warm water in the right ear. Suppose instead, you used cold water in the right ear. This causes this to be colder and go down. This comes up and clockwise movement of endolymph so you feel as if youre moving this way and you get left nystagmus. Thats two possibilities. The other possibilities are that you have the same guy lying on the same table and you do it in his left ear. And then finally, you turn the person face down and do it on both ears with warm and cold water. You have another four options. The easiest way to do is just figure out if head is back or forward, what the appropriate temperature is causing the endolymph to circulate this way or that way. The key here is that if someone seems to show function of his semicircular canals, dizziness, vertigo, etc then you might want to do this, both sides just to make sure that the vestibular systems are working properly to determine, narrow down the possibility where a lesion might exist. The other part of the vestibular system is the utricle. And there are two utricles: one on each side, roughly horizontal if you tilt your head forward. Utricle is bone covered. Youve got a surface called the macula. The macula has, like the crista of the semicircular canals, supportive endothelial cells, hair cells, and muscous secreting cells which puts a coating on the endothelial cells so that the cilia are embedded into the macula. What these organs have in addition is little stones, little crystals that embed itself in the gel that overlie the hair cells. That just gives them, the hair cells, this mucous layer, this gel, more mass. Youve got these calcium phosphate ittle crystals here floating around. They give it a little more mass, a little more weight and the thing is if you tilt your head, your macula is this way, the hair cells will be bent downhill. The other thing is if you look down on the macula youll see the hair cells and you can map them because each one has a longest cilium and a shortest one. What you find is therere all sorts of directions that the hair cells might be in over the whole surface of the macula so when you tilt it in one particular direction some will be depolarized and some will be hyperpolarized and some will be tilted sideways and not have any change at all. Now remember vestibular hair cells generate 5-10 AP/sec. so some will be hyperpolarized and their AP frequency will drop from 10 to 0 maybe. Whereas others will depolarize and their AP frequency will go up. And from the combination when there are two maculas, they are a mirror image of each other. So the corresponding place on this macula and the other one is depolarized and the other hyperpolarized. And what your vestibular nuclei do is get information about the distribution of APs coming from different parts of the nuclei of the utricle and interpret which way youre accelerating. Dont forget Einsteins theory of relativity. Gravity is the equivalent of acceleration. If you start moving this way the hair cells will be left behind and youll feel as though this way is down. Even though you are absolutely level. In regards to linear 6

Transcribed by _Jessica Li

February 16, 2014

acceleration, acceleration in a straight line, that you have to keep track of. A curiosity is that if you map out all the hair cells on the macula what youll find is that they are essentially aimed at an invisible nonexistent line. Which is called the striola. You wont see it but if you map the direction of the hair cells you will see that they are all aimed towards something. Is you see two go towards each other you just draw a line in-between, its an invisible, nonexistent line (the striola) because humans have a strange tendency to name things that dont exist. The striola gives you an idea of the orientation of the hair cells in the striola. And the point is if you tilt your head or accelerate, some hair cells depolarize, or hyperpolarize, or dont change t all. One other thing I mentioned a week ago when I was talking about sensory transducers I mentioned that the pacinian corpuscle in the utricle are very slow if not totally non adapting. Its not clear why.