12.4: Hemispheric Distribution
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After having dealt with how knowledge is stored in the brain, we now turn to the question of whether the brain is specialised and, if it is specialised, which functions are located where and which knowledge is present in which hemisphere. These questions can be subsumed under the topic “hemispheric specialisation” or “lateralisation of processing” which looks at the differences in processing between the two hemispheres of the human brain.
Differences between the hemispheres can be traced back to as long as 3.5 million years ago. Evidence for this are fossils of australopithecines (which is an ancient ancestor of homo sapiens). Because differences have been present for so long and survived the selective pressure they must be useful in some way for our cognitive processes.
Differences in Anatomy and Chemistry
Although at first glance the two hemispheres look identical, they differ in in various ways.
Concerning the anatomy, some areas are larger and the tissue contains more dendritic spines in one hemisphere than in the other. An example of this is what used to be called “Broca’s area” in the left hemisphere. This area which is –among other things- important for speech production shows greater branching in the left hemisphere than in the respective right hemisphere area. Because of the left hemisphere’s importance for language, with which we will deal later, one can conclude that anatomical differences have consequences for lateralisation in function.
Neurochemistry is another domain the hemispheres differ in: The left hemisphere is dominated by the neurotransmitter dopamine, whereas the right hemisphere shows higher concentrations of norepinephrine. Theories suggest that modules specialised on cognitive processes are distributed over the brain according to the neurotransmitter needed. Thus, a cognitive function relying on dopamine would be located in the left hemisphere.
The Corpus Callosum
The two hemispheres are interconnected via the corpus callosum, the major cortical connection. With its 250 million nerve fibres it is like an Autobahn for neural data connecting the two hemispheres. There are in fact smaller connections between the hemispheres but these are little paths in comparison. All detailed higher order information must pass through the corpus callosum when being transferred from one hemisphere to the other. The transfer time, which can be measured with ERP, lies between 5 and 20 ms.
Historic Approaches
Hemispheric specialisation has been of interest since the days of Paul Broca and Karl Wernicke, who discovered the importance of the left hemisphere for speech in the 1860s. Broca examined a number of patients who could not produce speech but whose understanding of language was not severed, whereas Wernicke examined patients who suffered the opposite symptoms (i.e. who could produce speech but did not understand anything). Both Broca and Wernicke found that their patients’ brains had damage to distinct areas of the left hemisphere.
Because in these days language was seen as the cognitive process superior to all other processes, the left hemisphere was believed to be superior to the right which was expressed in the “cerebral dominance theory” developed by J.H. Jackson. The right hemisphere was seen as a “spare tire [...] having few functions of its own” (Banich, S.94). This view was not challenged until the 1930s. In this decade and the following, research dramatically changed this picture. Of special importance for showing the role of the right hemisphere was Sperry, who conducted several experiments in 1974 for which he won the Nobel Prize in Medicine and Physiology in 1981.
Experiments with Split-Brain Patients
Sperry’s experiments took place with people who suffered a condition called “split brain syndrome” because they underwent a commissurotomy . In a commissurotomy the corpus callosum is sectioned so that communication between the hemispheres becomes severed in these patients. With his pioneering experiments, Sperry wanted to find out whether the left hemisphere really plays such an important role in speech processing as suggested by Broca and Wernicke.
Sperry used different experimental designs in his studies, but the basic assumption behind all experiments of this type was that perceptual information received at one side of the body is processed in the contra-lateral hemisphere of the brain. In one of the experiments the subjects had to recognise objects by touching it with merely one hand, while being blindfolded. He then asked the patients to name the object they felt and found that people could not name it when touching it with the left hand (which is linked to the right hemisphere). The question that arose was whether this inability was due to a possible function of the right hemisphere as “spare tire” or due to something else. Sperry now changed the design of his experiment so that patients now had to show that they recognised the objects by using it the right way. For example, if they recognised a pencil they would use it to write. With this changed design, no difference in performance between both hands were found.
In a different experiment conducted by Sperry et al. the patients were shown the word sky to one visual field and scraper to the other. They now had to draw the whole word they had seen with one hand. The patients were not able to synthesise this to skyscraper , instead they draw a scraper overlapped by some cloud. Thus it was concluded that each hemisphere took control of the hand to draw what it had seen.
Experiments with Patients with other Brain-Lesions
There have been other experiments conducted to gain more knowledge about hemispheric specialisation. They were conducted with epileptic individuals who were about to receive surgery where parts of one of their hemispheres was going to be removed. Before the surgery started it was important to find out which hemisphere is responsible for speech in this individual. This was done using the Wada-technique , where barbiturate is injected into one of the arteries supplying the brain with blood. Shortly after the injection, the contra-lateral side of the body is paralysed. If the person is now still able to speak, the doped hemisphere of the brain is not responsible for speech production in this individual. With the results of this technique it could be estimated that 95\% of all adult right-handers use their left hemisphere for speech.
Research with people who suffer brain lesions or even have a commissurotomy has some major draw backs: The reason why they had to undergo such surgery is usually epileptic seizures. Because of this, it is possible that their brains are not typical or have received damage to other areas during the surgery. Also, these studies have been performed with very limited numbers of subjects, so the statistical reliability might not be high.
Experiments with Neurologically Intact Individuals
In addition to experiments with brain-severed patients, studies with neurologically intact individuals have been conducted to measure perceptual asymmetries. These are usually performed with one of three methods: Namely the “divided visual field technique”, “dichaptic presentation” and “dichotic presentation”. Each of them again has as basic assumption the fact that perceptual information received at one side of the body is processed in the contra-lateral hemisphere.
Highly simplified picture of the visual pathway.
The divided visual field technique is based on the fact that the visual field can be divided into the right (RVF) and left visual field (LVF). Each visual field is processed independently from the other in the contra-lateral hemisphere. The divided visual field technique includes two different experimental designs: The experimenter can present one picture in just one of the visual fields and then let the subject respond to this stimulus. The other possibility involves showing two different pictures in each visual field.
A problem that can occur using the visual field technique is that the stimulus must be presented for less than 200 ms because this is how long the eyes can look at one point without shifting of the visual field.
In the dichaptic presentation technique the subject is presented two objects at the same time in each hand. (c.f. Sperry’s experiments)
The dichotic presentation technique enables researchers to study the processing of auditory information. Here, different information is presented simultaneously to each ear. Experiments with these techniques found that a sensory stimulus is processed 20 to 100 ms faster when it is initially directed to the specialised hemisphere for that task and the response is 10% more accurate.
Explanations for this include three hypotheses, namely the direct access theory , the callosal relay model and the activating-orienting model . The direct access theory assumes that information is processed in that hemisphere to which it is initially directed. This may result in less accurate responses, if the initial hemisphere is the unspecialised hemisphere. The Callosal relay model states that information if initially directed to the wrong hemisphere is transferred to the specialised hemisphere over the corpus callosum. This transfer is time-consuming and is the reason for loss of information during transfer. The activating-orienting model assumes that a given input activates the specialised hemisphere. This activation then places additional attention on the contra-lateral side of the activated hemisphere, “making perceptual information on that side even more salient”. (Banich)
Common Results
All the experiments mentioned above have some basic findings in common: The left hemisphere is superior at verbal tasks such as the processing of speech, speech production and recognition of letters whereas the right hemisphere excels at non-verbal tasks such as face recognition or tasks that involve spatial skills such as line orientation, or distinguishing different pitches of sound. This is evidence against the cerebral dominance theory which appointed the right hemisphere to be a spare tire! In fact both hemispheres are distinct and outclass at different tasks, and neither one can be omitted without this having high impact on cognitive performance.
Although the hemispheres are so distinct and are experts at their assigned functions, they also have limited abilities in performing the tasks for which the other hemisphere is specialised. In the picture above is an overview which hemisphere gives raise to what ability.
Differences in Processing
Experiment on local and global processing with patients with left- or right-hemisphere damage
There are two sets of approaches to the question of hemispheric specialisation. One set of theories is about the topic by asking the question “What tasks is each hemisphere specialised for?”. Theories that belong to this set, assign the different levels of ability to process sensory information to the different levels of abilities for higher cognitive skills. One theory that belongs to this set is the “spatial frequency hypothesis”. This hypothesis states that the left hemisphere is important for fine detail analysis and high spatial frequency in visual images whereas the right hemisphere is important for low spatial frequency. We have pursued this approach above.
The other approach does not focus on what type of information is processed by each hemisphere but rather on how each hemisphere processes information. This set of theories assumes that the left hemisphere processes information in an analytic, detail- and function-focused way and that it places more importance on temporal relations between information, whereas the right hemisphere is believed to go about the processing of information in a holistic way, focusing on spatial relations and on appearance rather than on function.
The picture above shows an exemplary response to different target stimuli in an experiment on global and local processing with patients who suffer right- or left-hemisphere damage. Patients with damage to the right hemisphere often suffer a lack of attention to the global form, but recognise details with no problem. For patients with left-hemisphere-damage this is true the other way around. This experiment supports the assumption that the hemispheres differ in the way they process information.
Interaction of the Hemispheres
Why is the transfer between the hemispheres needed at all if the hemispheres are so distinct concerning functioning, anatomy, chemistry and the transfer results in degrading of quality of information and takes time? The reason is that the hemispheres, although so different, do interact. This interaction has important advantages because as studies by Banich and Belger have shown it may “enhance the overall processing capacity under high demand conditions” (Banich). (Under low demand conditions the transfer does not make as much sense because the cost of transferring the information to the other hemisphere are higher than the advantages of parallel processing.)
The two hemispheres can interact over the corpus callosum in different ways. This is measured by first computing performance of each hemisphere individually and then measuring the overall performance of the whole brain. In some tasks one hemisphere may dominate the other in the overall performance, so the overall performance is as good or bad as the performance of one of the single hemispheres. What's surprising is that the dominating hemisphere may very well be the one that is less specialised, so here is another example of a situation where parallel processing is less effective than processing in just one half of the brain.
Another way of how the hemispheres interact is that overall processing is an average of performance of the two individual hemispheres.
The third, most surprising way the hemispheres can interact is that when performing a task together the hemispheres behave totally different than when performing the same task individually. This can be compared to social behavior of people: Individuals behave different in groups than they would when being by themselves.
Individual Factors Influencing Lateralisation
After having looked at hemispheric specialisation from a general point of view, we now want to focus on differences between individuals concerning hemispheric specialisation. Aspects that may have an impact on lateralisation might be age, gender or handed-ness.
Age could be one factor which decides in how far each hemisphere is used at specific tasks. Researchers have suggested that lateralisation develops with age until puberty. Thus infants should not have functionally-lateralised brains. Here are four pieces of evidence that speak against this hypothesis:
Infants already show the same brain anatomy as adults. This means the brain of a new born is already lateralised. Following the hypothesis that anatomy is linked to function this means that lateralisation is not developed at a later period in life.
Differences in perceptual asymmetries that means superior performance at processing verbal vs. non- verbal material in the different hemispheres cannot be observed in children aged 5 to 13, i.e. children aged 5 process the material the same way 13 year olds do.
Experiments with 1-week-old infants showed that they responded with increased interest to verbal material when this was presented to the right ear than when presented to the left ear and increased interest to non-verbal material when presented to the left ear. The infants’ interest was hereby measured by the frequency of soother sucking.
Although children who underwent hemispherectomy (the surgical removal of one hemisphere) do develop the cognitive skills of the missing hemisphere (in contrast to adults or adolescents who can only partly compensate for missing brain parts), they do not develop these skills to the same extent as a child with hemispherectomy of the other hemisphere. For example: A child whose right hemisphere has been removed will develop spatial skills but not to the extent that a child whose left hemisphere has been removed, and thus still possesses the right hemisphere.
Handedness is another factor that might influence brain lateralisation. There is statistical evidence that left-handers have a different brain organisation than right-handers. 10% of the population is left-handed. Whereas 95% of the right-handed people process verbal material in a superior manner in the left-hemisphere, there is no such a high figure for verbal superiority of one hemisphere in left-handers: 70% of the left-handers process verbal material in the left-hemisphere, 15% process verbal material in the right hemisphere (the functions of the hemispheres are simply switched around), and the remaining 15% are not lateralised, meaning that they process language in both hemispheres. Thus as a group, left-handers seem to be less lateralised. However a single left-handed-individual can be just as lateralised as the average right-hander.
Gender is also an aspect that is believed to have impact on the hemispheric specialisation. In animal studies, it was found that hormones create brain differences between the genders that are related to reproductional functions. In humans it is hard to determine to which extent it is really hormones that cause differences and to which extent it is culture and schooling that are responsible.
One brain area for which a difference between the genders was observed is the corpus callosum. Although one study found that the c.c. is larger in women than in men these results could not be replicated. Instead it was found that the posterior part of the c.c. is more bulbous in women than in men. This might however be related to the fact that the average woman has a smaller brain than the average man and thus the bulbousness of the posterior section of the c.c. might be related to brain size and not to gender.
In experiments that measure performance in various tasks between the genders the cultural aspect is of great importance because men and women might use different problem solving strategies due to schooling.
Summary
Although the two hemispheres look like each other’s mirror images at first glance, this impression is misleading. Taking a closer look, the hemispheres not only differ in their conformation and chemistry, but most importantly in their function. Although both hemispheres can perform all basic cognitive tasks, there exists a specialisation for specific cognitive demands. In most people, the left hemisphere is an expert at verbal tasks, whereas the right hemisphere has superior abilities in non-verbal tasks. Despite the functional distinctness the hemispheres communicate with each other via the corpus callosum.
This fact has been utilised by Sperry’s experiments with split-brain-patients. These are outstanding among other experiments measuring perceptual asymmetries because they were the first experiments to refute the hemispheric dominance theory and received recognition through the Nobel Prize for Medicine and Physiology.
Individual factors such as age, gender or handed-ness have no or very little impact on hemispheric functioning.