The cerebellum is located to the brainstem and to the occipital lobe of the cerebrum
5.1 Overview: Functions of the Cerebellum Show The cerebellum (“little brain”) is a structure that is located at the back of the brain, underlying the occipital and temporal lobes of the cerebral cortex (Figure 5.1). Although the cerebellum accounts for approximately 10% of the brain’s volume, it contains over 50% of the total number of neurons in the brain. Historically, the cerebellum has been considered a motor structure, because cerebellar damage leads to impairments in motor control and posture and because the majority of the cerebellum’s outputs are to parts of the motor system. Motor commands are not initiated in the cerebellum; rather, the cerebellum modifies the motor commands of the descending pathways to make movements more adaptive and accurate. The cerebellum is involved in the following functions: Maintenance of balance and posture. The cerebellum is important for making postural adjustments in order to maintain balance. Through its input from vestibular receptors and proprioceptors, it modulates commands to motor neurons to compensate for shifts in body position or changes in load upon muscles. Patients with cerebellar damage suffer from balance disorders, and they often develop stereotyped postural strategies to compensate for this problem (e.g., a wide-based stance). Coordination of voluntary movements. Most movements are composed of a number of different muscle groups acting together in a temporally coordinated fashion. One major function of the cerebellum is to coordinate the timing and force of these different muscle groups to produce fluid limb or body movements. Motor learning. The cerebellum is important for motor learning. The cerebellum plays a major role in adapting and fine-tuning motor programs to make accurate movements through a trial-and-error process (e.g., learning to hit a baseball). Cognitive functions. Although the cerebellum is most understood in terms of its contributions to motor control, it is also involved in certain cognitive functions, such as language. Thus, like the basal ganglia, the cerebellum is historically considered as part of the motor system, but its functions extend beyond motor control in ways that are not yet well understood. 5.2 Cerebellar Gross Anatomy The cerebellum consists of two major parts (Figure 5.2A). The cerebellar deep nuclei (or cerebellar nuclei) are the sole output structures of the cerebellum. These nuclei are encased by a highly convoluted sheet of tissue called the cerebellar cortex, which contains almost all of the neurons in the cerebellum. A cross-section through the cerebellum reveals the intricate pattern of folds and fissures that characterize the cerebellar cortex (Figure 5.3). Like the cerebral cortex, cerebellar gyri are reproducible across individuals and have been identified and named. We will only be concerned with some of the larger divisions of the cerebellar cortex.
Divisions of the cerebellum. Two major fissures running mediolaterally divide the cerebellar cortex into three primary subdivisions (Figure 5.2B and Figure 5.3). The posterolateral fissure separates the flocculonodular lobe from the corpus cerebelli, and the primary fissure separates the corpus cerebelli into a posterior lobe and an anterior lobe (Figure 5.4). The cerebellum is also divided
sagittally into three zones that run from medial to lateral (Fig. 5.4). The vermis (from the Latin word for worm) is located along the midsagittal plane of the cerebellum. Directly lateral to the vermis is the intermediate zone. Finally, the lateral hemispheres are located lateral to the intermediate zone (there are no clear morphological borders between the intermediate zone and the lateral hemisphere that are visible from a gross
specimen).
Cerebellar nuclei. All outputs from the cerebellum originate from the cerebellar deep nuclei. Thus, a lesion to the cerebellar nuclei has the same effect as a complete lesion of the entire cerebellum. It is important to know the inputs, outputs, and anatomical relationships between the different cerebellar nuclei and the subdivisions of the cerebellum (Figure 5.5).
In addition to these inputs, all cerebellar nuclei and all regions of cerebellum get special inputs from the inferior olive of the medulla (discussed below). It is convenient to remember that the anatomical locations of the cerebellar nuclei correspond to the cerebellar cortex regions from which they receive input. Thus, the medially located fastigial nucleus receives input from the medially located vermis; the slightly lateral interposed nuclei receive input from the slightly lateral intermediate zone; and the most lateral dentate nucleus receives input from the lateral hemispheres. Cerebellar peduncles. Three fiber bundles carry the input and output of the cerebellum.
Thus, the inputs to the cerebellum are conveyed primarily through the inferior and middle cerebellar peduncles, whereas the outputs are conveyed primarily through the superior cerebellar peduncle. The inputs arise from the ipsilateral side of the body, and the outputs also go to the ipsilateral side of the body. Note that this is true even for the outputs to the contralateral red nucleus. Recall from the chapter on descending motor pathways that the rubrospinal tract immediately crosses the midline after the fibers leave the red nucleus. Thus, cerebellar output to the red nucleus affects the ipsilateral side of the body by a double-crossed pathway. Unlike the cerebral cortex, the cerebellum receives input from, and controls output to, the ipsilateral side of the body, and damage to the cerebellum therefore results in deficits to the ipsilateral side of the body. 5.3 Functional Subdivisions of the Cerebellum The anatomical subdivisions described above correspond to three major functional subdivisions of the cerebellum. Vestibulocerebellum. The vestibulocerebellum comprises the flocculonodular lobe and its connections with the lateral vestibular nuclei. Phylogenetically, the vestibulocerebellum is the oldest part of the cerebellum. As its name implies, it is involved in vestibular reflexes (such as the vestibuloocular reflex; see below) and in postural maintenance. Spinocerebellum. The spinocerebellum comprises the vermis and the intermediate zones of the cerebellar cortex, as well as the fastigial and interposed nuclei. As its name implies, it receives major inputs from the spinocerebellar tract. Its output projects to rubrospinal, vestibulospinal, and reticulospinal tracts. It is involved in the integration of sensory input with motor commands to produce adaptive motor coordination. Cerebrocerebellum. The cerebrocerebellum is the largest functional subdivision of the human cerebellum, comprising the lateral hemispheres and the dentate nuclei. Its name derives from its extensive connections with the cerebral cortex, via the pontine nuclei (afferents) and the VL thalamus (efferents). It is involved in the planning and timing of movements. In addition, the cerebrocerebellum is involved in the cognitive functions of the cerebellum. 5.4 Histology and Connectivity of Cerebellar Cortex The cerebellar cortex is divided into three layers (Figure 5.6). The innermost layer, the granule cell layer, is made of 5 x 1010 small, tightly packed granule cells. The middle layer, the Purkinje cell layer, is only 1-cell thick. The outer layer, the molecular layer, is made of the axons of granule cells and the dendrites of Purkinje cells, as well as a few other cell types. The Purkinje cell layer forms the border between the granule and molecular layers.
Granule cells. Granule cells are very small, densely packed neurons that account for the huge majority of neurons in the cerebellum. Indeed, cerebellar granule cells account for more than half of the neurons in the entire brain. These cells receive input from mossy fibers and project to the Purkinje cells.
Purkinje cells. The Purkinje cell is one of the most striking cell types in the mammalian brain. Its apical dendrites form a large fan of finely branched processes (Figure 5.7). Remarkably, this dendritic tree is almost two-dimensional; looked at from the side, the dendritic tree is flat (click PLAY on Figure 5.7). Moreover, all Purkinje cells are oriented in parallel. This arrangement has important functional considerations, as we shall see below. Connectivity. The cerebellar cortex has a relatively simple, stereotyped connectivity pattern that is identical throughout the whole structure. Figure 6 illustrates a simplified diagram of the connectivity of the cerebellum. Cerebellar input can be divided into two distinct classes.
The Purkinje cell is the sole source of output from the cerebellar cortex. It is important to note that Purkinje cells make inhibitory connections onto the cerebellar nuclei. (Note the distinction between the Purkinje cells, which constitute the sole output of the cerebellar cortex, and the cerebellar nuclei, which constitute the sole output of the entire cerebellum.) Almost all of the spikes generated by the Purkinje cell are caused by its parallel-fiber inputs. These inputs cause the Purkinje cell to fire at a high resting rate (~70 spikes/sec), tonically inhibiting its cerebellar nucleus targets. The powerful inputs from climbing fibers occur less frequently (~1 spike/sec); thus, they have a minor influence on the overall firing rate of the Purkinje cell. The Purkinje cell spikes that are generated by climbing fibers are calcium-spikes, however, which allow the climbing fibers to initiate a number of calcium-dependent changes in the Purkinje cell. As described below, one important change appears to be a long-lasting change in the strength of the parallel-fiber inputs to the Purkinje cell. 5.5 Damage to Cerebellum Produces Movement Disorders Much of what is known about cerebellar function comes from studies of patients with cerebellar damage. In general, such patients display uncoordinated voluntary movements and problems maintaining balance and posture. The following are some symptoms of cerebellar damage (we will discuss more symptoms in the next chapter):
A second example of cerebellum-dependent motor learning involves the execution of accurate, coordinated movements. Subjects wore prism goggles that shifted the visual image to the right, and they were asked to then throw balls at a target on the wall. Because of the prisms, the accuracy of the subjects was initially quite low, as the balls consistently hit to the left of the target. With repeated practice, however, the subjects became more and more accurate at hitting the target. When the goggles were removed, the subject now began to throw the balls to the right of the target, because their motor programs had been recalibrated to use the shifted visual input. Over time, once again, they gradually increased their accuracy. Patients with cerebellar damage never learned to compensate for the prism, as their balls always landed to the left of the target when the goggles were worn. When the goggles were removed, they were immediately accurate at hitting the target, because they never made compensations for the earlier prism trials. A third example involves the Pavlovian classical conditioning of the eye blink reflex. In this task, a neutral stimulus (such as a tone) is paired with a noxious stimulus (such as a puff of air to the eye) that causes a reflexive eye blink. Over time, experimental animals will learn to close their eye when the tone occurs, in anticipation of the air puff. This learned eyelid closure is remarkably well-timed to peak at the expected time of the puff. Animals with cerebellar damage do not learn to produce the eyelid closure in response to the tone. 5.6 Cerebellum and Control Systems What do the various symptoms of cerebellar damage have in common that reveal the function of the cerebellum? A number of different theories have been proposed. Recall the discussion in Chapter 1 of the ubiquitous use of sensory information in motor control. The cerebellum receives extensive sensory input, and it appears to use this input to guide movements in both a feedback and feedforward control manner. Feedback control systems In a feedback controller, a desired output is compared continuously with the actual output, and adjustments are made during the execution of the movement until the actual movement matches the desired movement. A common example of a feedback control system is the thermostat in your home (Figure 5.9).
The thermostat is set to a desired temperature (e.g., 72°), and a thermometer measures the current temperature in the room. If the thermostat (the comparator) detects that the room is cooler than the desired temperature, it sends an error signal that turns on the furnace. If the comparator detects that the room is warmer than the desired setting, its sends an error signal that turns on the air conditioner. Feedback control systems can produce very accurate outputs; however, in general they are slow. In order to change the output, the effector must wait until information is transmitted from the sensor to the comparator and then to the effector. At this point, another comparison is made, and the process continues. Consider further the thermostat example. If the temperature reads 5° cooler than desired, the thermostat can instruct the furnace to turn on at a moderate heat. It reads the new room temperature, and, if it is still too cool, it instructs the furnace to deliver more heat, and so on. Although this will eventually produce an accurate room temperature at the desired point, it takes a number of cycles to reach that point. One possible solution for quicker results would be to turn an enormous furnace on full-blast, such that is heats the room very quickly. This solution, however, can generate another problem. It will tend to cause the system to oscillate if the feedback pathways are slow. For example, assume that the furnace can heat the room at the rate of 5° per second, but that it takes 2 seconds for the thermometer to adjust to the new temperature, and for the new error signal to turn the furnace off. In those 2 seconds, the furnace has heated the room up 10°, and now it is too warm. So the error signal turns on the air conditioner, and it cools the room at 5°/sec. Of course, it also takes 2 sec to receive the feedback, and by the time it is told to shut off, it has cooled the room by 10°. You can see what happens: the system will be sent into an endless oscillation of being 5° too hot and 5° too cold. In order for a feedback system to work well, the transmission time of sensory information through the comparator to the effector must be rapid compared to the time of the action. Feedback control systems work well only when the sensory feedback about the actual output is fast relative to the actual output. If the actual output is faster than the sensor’s ability to provide feedback, then the system will tend to oscillate between overshooting and undershooting the desired output. Thus, a feedback controller is useful for slow movements, like postural adjustments. The role of the myotatic reflex in posture maintenance is an example of a feedback controller in the spinal cord, and the cerebellum plays a role in coordinating these postural adjustments. Feedback control is not effective for most of the fast movements we make routinely (such as an eye movement or reaching out for a cup). For these movements, a feedforward controller is needed. Feed forward control systems In a feedforward control system, when a desired output is sent to the controller, the controller evaluates sensory information about the environment and about the system itself before the output commands are generated. It uses the sensory information to program the best set of instructions to accomplish the desired output. However, in a pure feedforward system, once the commands are sent, there is no way to alter them (i.e., there is no feedback loop). The advantage of a feedforward system is that it can produce the precise set of commands for the effector without needing to constantly check the output and make corrections during the movement itself. The main disadvantage, however, is that the feedforward controller requires a period of trial-and-error learning before it can function properly. In most biological systems, it is hard (perhaps impossible) to pre-program all of the possible sensory conditions that the controller may encounter during the life of the organism. Furthermore, the environment and conditions under which actions are made are constantly changing, and the feedforward controller must be able to adapt its output commands to account for these changes.
Let us extend the thermostat example to see how a temperature controller operating as a feedforward system would work to raise the temperature of a room from 70° to 75°. The controller would use diverse sensory information about the environment before sending its command to the furnace (Figure 5.10). For example, it would read the current temperature, the current humidity level, the size of the room, the number of people in the room, and so forth. Based on this information, it would direct the furnace to turn on for a pre-set period of time, and that’s it. There would be no need to continually compare the current temperature with the desired setting, as the system has predetermined how long the furnace needs to be working in order to achieve the desired temperature. How did the controller obtain this information? A feedforward controller requires a large amount of experience in order to learn the appropriate actions needed for each set of environmental conditions. If on one trial it turns the furnace off too soon and the room does not reach the desired temperature, it adjusts its programming such that the next time it encounters the same environmental conditions, it turns the furnace on for a longer period of time. Through many such instances of trial and error learning, the feedforward system creates a “look-up table” that tells it how long the furnace needs to be active under the current conditions. The key distinction between a feedback and feedforward system is that the feedback system uses sensory information to generate an error signal during the control of a movement, whereas a feedforward system uses sensory information in advance of a movement. Any error signal about the final output is used by the feedforward system only to change its programming of future movements. The cerebellum may be a feedforward control system The cerebellar involvement in the VOR may be explained in terms of the learning requirements of a feedforward controller. When the head moves, a compensatory eye movement must be made to maintain a stable gaze. The cerebellum receives sensory input from the vestibular system informing it that the head is moving. It also receives input from eye muscle proprioceptors and other relevant sources of information about current conditions in order to make an accurate compensatory eye movement. It evaluates all of this advance sensory information and calculates the proper eye movement to exactly counterbalance the head movement. What if the eye movement does not match the head movement, however, and the visual image moves across the retina (such as in the experimental condition in which a prism was worn, or in a real-life situation in which an individual wears new prescription eyeglasses)? The retinal slip constitutes an error signal to tell the cerebellum that next time these conditions are met, adjust the eye movement to decrease the retinal slip. This trial and error sequence will be repeated until the movement is properly calibrated; moreover, these mechanisms will ensure that the movements stay calibrated. As another example, the coordination of movements requires that muscle groups be activated in precise temporal sequence. Not only do the different joints need to be coordinated temporally, but even antagonist muscles that control the same joint need precise temporal coordination. For example, an extensor muscle needs to be activated to
start a reaching movement, and the corresponding flexor muscle needs to be activated at the end of the movement to stop the movement appropriately. The precise timing of muscle contractions and the force necessary for each contraction varies with the amount of load placed on a muscle, as well as on the inherent properties of the muscle itself (e.g., elasticity). These variables are constantly changing throughout life, as one grows, gains/loses weights, and ages. Moreover, a similar movement will
require different patterns of motor activity depending on the weight being born by the muscle (for example, if an extended hand is empty or holding a heavy weight). The cerebellum appears necessary for the proper timing and coordination of muscle groups, very likely through a trial-and-error learning mechanism discussed previously. Such a role helps explain the deficits seen in dysdiadochokinesia, in which patients cannot perform rapidly alternating sequences of movements.
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The spinocerebellum contains the...
The spinocerebellum contains the...
The spinocerebellum contains the...
The spinocerebellum contains the...
The spinocerebellum contains the...
The spinocerebellum contains the...
The lateral vestibular nuclei are functionally analogous to the...
The lateral vestibular nuclei are functionally analogous to the...
The lateral vestibular nuclei are functionally analogous to the...
The lateral vestibular nuclei are functionally analogous to the...
The lateral vestibular nuclei are functionally analogous to the...
The lateral vestibular nuclei are functionally analogous to the...
Donations to Neuroscience Online will help fund development of new features and content. Where is the cerebrum and cerebellum located?There are three major components of the brain. The cerebrum is the largest component, extending across the top of the head down to ear level. The cerebellum is smaller than the cerebrum and located underneath it, behind the ears toward the back of the head.
Where is your cerebellum located?Your cerebellum is a part of your brain located at the back of your head, just above and behind where your spinal cord connects to your brain itself. The name “cerebellum” comes from Latin and means “little brain.” For centuries, scientists believed your cerebellum's job was to coordinate your muscle movements.
Is the cerebellum inferior to the occipital and temporal lobes?The cerebellum lies inferior to the occipital lobes. The cerebellum is also divided into two hemispheres, like the cerebral cortex. The cerebellum is best known for its role in regulation and control of movement, but it is also involved in cognitive functions like emotions.
Is the cerebellum anterior or posterior to the brainstem?Definition: A part of the central nervous system found posteriorly to the brainstem that is in charge for motor learning, coordination and precision of motor functions.
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