The neurons of the cerebral cortex constitute the highest level of control in the hierarchy of the nervous system. Consequently, the terms higher cerebral functions and higher cortical functions are used by neurologists and neuroscientists to refer to all conscious mental activity, such as thinking, remembering, and reasoning, and to complex volitional behaviour such as speaking and carrying out purposive movement. The terms also refer to the processing of information in the cerebral cortex, most of which takes place unconsciously.
Neuroscientists investigate the structure and functions of the cerebral cortex, but the processes involved in thinking are also studied by cognitive psychologists, who group the mental activities known to the neuroscientist as higher cortical functions under the headings cognitive function or human information processing. From this perspective, complex information processing is the hallmark of cognitive function. Cognitive science attempts to identify and define the processes involved in thinking without regard to their physiological basis. The resulting models of cognitive function resemble flowcharts for a computer program more than neural networks—and, indeed, they frequently make use of computer terminology and analogies.
The discipline of neuropsychology, by studying the relationship between behaviour and brain function, bridges the gap between neural and cognitive science. Examples of this bridging role include studies in which cognitive models are used as conceptual frameworks to help explain the behaviour of patients who have suffered damage to different parts of the brain. Thus, damage to the frontal lobes can be conceptualized as a failure of the “central executive” component of working memory, and a failure of the “generate” function in another model of mental imagery would fit with some of the consequences of left parietal lobe damage.
The analysis of changes in behaviour and ability following damage to the brain is by far the oldest and probably the most-informative method adopted for studying higher cortical functions. Usually these changes take the form of what is known as a deficit—that is, an impairment of the ability to act or think in some way. With certain stipulations, one can assume that the damaged part of the brain is involved in the function that has been lost. However, people vary considerably in their abilities, and most brain lesions occur in subjects whose behaviour was not formally studied before they became ill. Lesions are rarely precisely congruent with the brain area responsible for a given function, and their exact location and extent can be difficult to determine even with modern imaging techniques. Abnormal behaviour after brain injury, therefore, is often difficult to attribute to precisely defined damage or dysfunction.
It would also be naive to suppose that a function is represented in a particular brain area just because it is disrupted after damage to that area. For example, a tennis champion does not play well with a broken ankle, but this would not lead one to conclude that the ankle is the centre in which athletic skill resides. Reasonably certain conclusions about brain-behaviour relationships, therefore, can be drawn only if similar well-defined changes occur reliably in a substantial number of patients suffering from similar lesions or disease states.
The most prominent series of observations clearly belonging to modern neuropsychology were made by Paul Broca in the 1860s. He reported the cases of several patients whose speech had been affected following damage to the left frontal lobe and provided autopsy evidence of the location of the lesion. Broca explicitly recognized the left hemisphere’s control of language, one of the fundamental phenomena of higher cortical function.
In 1874 the German neurologist Carl Wernicke described a case in which a lesion in a different part of the left hemisphere, the posterior temporal region, affected language in a different way. In contrast to Broca’s cases, language comprehension was more affected than language output. This meant that two different aspects of higher cortical function had been found to be localized in different parts of the brain. In the next few decades there was a rapid expansion in the number of cognitive processes studied and tentatively localized.
Wernicke was one of the first to recognize the importance of the interaction between connected brain areas and to view higher cortical function as the buildup of complex mental processes through the coordinated activities of local regions dealing with relatively simple, predominantly sensory-motor functions. In doing so, he opposed the view of the brain as an equipotential organ acting en masse.
Since Wernicke’s time, scientific views have swung between the localization and mass-action theories. Major advances in the 20th century included vast increases in knowledge, the discovery of new ways of studying the anatomy and physiology of the brain, and the introduction of better quantitative methods in the study of behaviour.
Broca’s declaration that the left hemisphere is predominantly responsible for language-related behaviour is the clearest and most dramatic example of an asymmetry of function in the human brain. This functional asymmetry is related to hand preference and probably to anatomical differences, although neither relationship is simple.
Evidence from a number of converging sources, notably the high incidence of the language disturbance aphasia after left- but not right-hemisphere damage, indicates that the left hemisphere is dominant for the comprehension and expression of language in close to 99 percent of right-handed people. At least 60 percent of left-handed and ambidextrous people also have left-hemisphere language, but up to 30 percent have predominantly right-hemisphere language. The remainder have language represented to some degree in both hemispheres.
The posterior temporal region of the brain, which is one of the regions responsible for language in the dominant hemisphere, is physically asymmetrical; specifically, the area known as the planum temporale is larger in the left hemisphere in most people. This asymmetry is more common in right-handers, while left-handed individuals are likely to have more nearly symmetrical brains. Reduced anatomical asymmetry has also been found in people with right-hemisphere dominance for speech and in some people with the reading disorder dyslexia. These results point to some relationship between handedness, cerebral dominance for language, anatomical asymmetry in the temporal lobe, and some aspects of language competence. Certainly there is a tendency for right-handedness, left-hemisphere dominance for language, and a larger left planum temporale to occur together. However, there are exceptions; for example, a few right-handers are right-hemisphere dominant for speech, and some right-handers who have left-hemisphere speech do not have a larger left planum temporale. In people who are atypical in one of these respects—for example, by being left-handed—the relationship between handedness, cerebral dominance, and anatomical asymmetry is much less consistent. Therefore, language is not invariably located in the hemisphere opposite the dominant hand or in the hemisphere with the larger planum temporale.
Studies of individuals being treated for epilepsy in whom the corpus callosum (the bundle of nerve fibres connecting the two halves of the brain) has been severed, allowing the two hemispheres to function largely independently, have revealed that the right hemisphere has more language competence than was thought. These individuals show evidence of comprehension of words presented to the isolated right hemisphere, although that hemisphere is not able to initiate speech. The speech of individuals with a lesion of the right hemisphere may lack normal melodic quality, and they may have difficulty expressing and understanding such things as emotional overtones. They may also have difficulty appreciating some of the more subtle, connotative aspects of language, such as puns, figures of speech, and jokes. Nevertheless, the dominance of the left hemisphere for language, particularly the syntactic aspects of language and language output, is the clearest example yet discovered of the lateralization of higher cortical function.
The left hemisphere also appears to be more involved than the right in the programming of complex sequences of movement and in some aspects of awareness of one’s own body. Thus, apraxia is more common after damage to the left hemisphere. In apraxia, the individual has difficulty performing actions involving several movements or the manipulation of objects in an appropriate and skillful way. The difficulty appears to be in programming the motor system to control the sequence of movements required to perform a complex action in the appropriate order and with the appropriate timing.
Confusion of right and left is also found after left-hemisphere damage, making it appear that the left hemisphere is largely responsible for collating somatosensory information into a special awareness of the body called the body image. Finger agnosia is a condition in which the individual does not appear to “know” which finger is which and is unable to indicate which one the examiner touches without the aid of vision. The phenomenon of the phantom limb, whereby patients “feel” sensations in amputated limbs, indicates that the brain’s internal representation of the body may persist intact for some time after the loss of a body part. This internal representation appears to be maintained chiefly by the left hemisphere.
The special functions of the right hemisphere were recognized later than those of the left hemisphere, although a case of “imperception” reported by the English neurologist John Hughlings Jackson in 1876 foreshadowed later findings. Jackson’s patient, who had a lesion in the posterior part of the right hemisphere, lost her way in familiar surroundings, failed to recognize familiar places and people, and had difficulty in dressing herself—all of which became well-recognized consequences of right-hemisphere damage. The right hemisphere appears to be specialized for some aspects of higher-level visual perception, spatial orientation, and sense of direction, and it probably plays a dominant role in the recognition of objects and faces. The specialization of the right hemisphere, however, is less absolute than that of the left hemisphere in that these skills are less lateralized than language.
There has been considerable speculation as to why the human brain is functionally asymmetrical. Initially, both functional and anatomical asymmetry were thought, like language, to be a uniquely human trait, but less-pronounced asymmetries have now been found in lower animals. One theory is that it is necessary to have language represented in a single hemisphere to avoid competition between the hemispheres for control of the muscles involved in speech. Another theory is that it is efficient to have the language system represented in a restricted area on one side of the brain because information needs to be transferred over short distances and fewer connections. A third theory is that the dominance of the left hemisphere over the right hand and skilled movement preceded its dominance over language. According to this view, language subsequently developed in the same hemisphere because language implies speech, which requires precise programming of sequences of movement in the articulatory musculature. None of these theories has been conclusively proved correct or has been generally accepted. Also, there remain some facts that are difficult to explain by any theory. For example, all of the above theories would predict that bilateral and, in some cases, right-hemisphere language representation would be disadvantageous, but this does not seem to be generally true.