Even in fish, lesions of the forebrain destroy the traits of initiative and caution. In higher animals these traits, much elaborated, seem localized in the neo-cortex, the site of many of the characteristic human cognitive functions. It is frequently discussed in terms of four major regions or lobes: the frontal, parietal, temporal and occipital lobes. Early neurophysiologists held that the neocortex was primarily connected only to other places in the neocortex, but it is now Known that there are many neural connections with the sub-cortical brain. It is, however, by no means clear that the neocortical subdivisions are actually functional units. Each certainly has many quite different functions, and some functions may be shared among or between lobes. Among other functions, the frontal lobes seem to be connected with deliberation and the regulation of action; the parietal lobes, with spatial perception and the exchange of information between the brain and the rest of the body; the temporal lobes, with a variety of complex perceptual tasks; and the occipital lobes, with vision, the dominant sense in humans and other primates.
For many decades the prevailing view of neurophysiologists was that the frontal lobes, behind the forehead, are the sites of anticipation and planning for the future, both characteristically human functions. But more recent work has shown that the situation is not so simple. A large number of cases of frontal lesions-largely suffered in warfare and as gunshot wounds-have been investigated by the American neurophysiologist Hans-Lukas Teuber of the Massachusetts Institute of Technology. He found that many frontal-lobe lesions have almost no obvious effects on behavior; however, in severe pathology of the frontal lobes "the patient is not altogether devoid of capacity to anticipate a course of events, but cannot picture himself in relation to those events as a potential agent." Teuber emphasized the fact that the frontal lobe may be involved in motor as well as cognitive anticipation, particularly in estimating what the effect of voluntary movements will be. The frontal lobes also seem to be implicated in the connection between vision and erect bipedal posture.
Thus the frontal lobes may be involved with peculiarly human functions in two different ways. If they control anticipation of the future, they must also be the sites of concern, the locales of worry. This is why transection of the frontal lobes reduces anxiety. But prefrontal lobotomy must also greatly reduce the patient's capacity to be human. The price we pay for anticipation of the future is anxiety about it. Foretelling disaster is probably not much fun; Pollyanna was much happier than Cassandra. But the Cassandric components of our nature are necessary for survival. The doctrines for regulating the future that they produced are the origins of ethics, magic, science and legal codes. The benefit of foreseeing catastrophe is the ability to take steps to avoid it, sacrificing short-term for long-term benefits. A society that is, as a result of such foresight, materially secure generates the leisure time necessary for social and technological innovation.
The other suspected function of the frontal lobes is to make possible mankind's bipedal posture. Our upright stance may not have been possible before the development of the frontal lobes. As we shall see later in more detail, standing on our own two feet freed our hands for manipulation, which then led to a major accretion of human cultural and physiological traits. In a very real sense, civilization may be a product of the frontal lobes.
Visual information from the eyes arrives in the human brain chiefly in the occipital lobe, in the back of the head; auditory impressions, in the upper part of the temporal lobe, beneath the temple. There is fragmentary evidence that these components of the neo-cortex are substantially less well developed in blind deaf-mutes. Lesions in the occipital lobe-as produced by gunshot wounds, for example-frequently induce an impairment in the field of vision. The victim may be in all other respects normal but able to see only with peripheral vision, perceiving a solid, dark blot looming in front of him at the center of the normal field of view. In other cases, more bizarre perceptions follow, including geometrically regular, cursive floating impairments in the visual field, and "visual fits" in which (for example) objects on the floor to the patient's lower right are momentarily perceived as floating in the air to his upper left and rotated 180 degrees through space. It may even be possible to map which parts of the occipital lobes are responsible for which visual functions by systematically calculating the impairments of vision from various occipital lesions. Permanent impairments of vision are much less likely to occur in the very young, whose brains seem able to repair themselves or transfer functions to neighboring regions very well.
The ability to connect auditory with visual stimuli is also localized in the temporal lobe. Lesions in the temporal lobe can result in a form of aphasia, the inability to recognize spoken words. It is remarkable and significant that brain-damaged patients can be completely competent in spoken language and entirely incompetent in written language, or vice versa. They may be able to write but unable to read; able to read numbers but not letters; able to name objects but not colors. There is in the neocortex a striking separation of function, which is contrary to such common-sense notions as that reading and writing, or recognizing words and numbers, are very similar activities. There are also as yet unconfirmed reports of brain damage that results only in the inability to understand the passive voice or prepositional phrases or possessive constructions. (Perhaps the locale of the subjunctive mood will one day be found. Will Latins turn out to be extravagantly endowed and English-speaking peoples significantly short-changed in this minor piece of brain anatomy?) Various abstractions, including the "parts of speech" in grammar, seem, astonishingly, to be wired into specific regions of the brain.
In one case, a temporal-lobe lesion resulted in a surprising impairment in the patient's perception of faces, even the faces of his immediate family. Presented with the face on this page, he described it as "possibly" being an apple. Asked to justify this interpretation, he identified the mouth as a cut in the apple, the nose as the stem of the apple folded back on its surface, and the eyes as two worm holes. The same patient was perfectly able to recognize sketches of houses and other inanimate objects. A wide range of experiments shows that lesions in the right temporal lobe produce amnesia for certain types of nonverbal material, while lesions in the left temporal lobe produce a characteristic loss of memory for language.
Our ability to read and make maps, to orient ourselves spatially in three dimensions and to use the appropriate symbols-all of which are probably involved in the development if not the use of language -are severely impaired by lesions in the parietal lobes, near the top of the head. One soldier who suffered a massive wartime penetration of the parietal lobe was for a full year unable to orient his feet into his slippers, much less find his bed in the hospital ward. He nevertheless eventually experienced an almost complete recovery. A lesion of the angular gyrus of the neocortex, in the parietal lobe, results in alexia, the inability to recognize the printed word. The parietal lobe appears to be involved in all human symbolic language and, of all the brain lesions, a lesion in the parietal lobe causes the greatest decline in intelligence as measured by activities in everyday life.
Chief among the neocortical abstractions are the human symbolic languages, particularly reading and writing and mathematics. These seem to require cooperative activities of the temporal, parietal and frontal lobes, and perhaps the occipital as well. Not all symbolic languages are neocortical however; bees- without a hint of a neocortex-have an elaborate dance language, first elucidated by the Austrian entomologist Karl von Frisch, by which they communicate information on the distance and direction of available food. It is an exaggerated gestural language, imitative of the motions bees in fact perform when finding food-as if we were to make a few steps towards the refrigerator, point and rub our bellies, with our tongues lolling out all the while. But the vocabularies of such languages are extremely limited, perhaps only a few dozen words. The kind of learning that human youngsters experience during their long childhood seems almost exclusively a neocortical function.
While most olfactory processing is in the limbic system, some occurs in the neocortex. The same division of function seems to apply to memory. A principal part of the limbic system, other than the olfactory cortex, is, as we have mentioned, the hippocampal cortex. When the olfactory cortex is excised, animals can still smell, although with a much lower efficiency. This is another demonstration of the redundancy of brain function. There is some evidence that, in contemporary humans, the short-term memory of smell resides in the hippocampus. The original function of the hippocampus may have been exclusively the short-term memory of smell, useful in, for example, tracking prey or finding the opposite sex. But a bilateral hippocampal lesion in humans results, > as in the case of H. M., in a profound impairment of all varieties of short-term memory. Patients with such lesions literally cannot remember from one moment to the next. Clearly, both hippocampus and frontal lobes are involved in human short-term memory.
One of the many interesting implications of this is that short-term and long-term memory reside mostly in different parts of the brain. Classical conditioning- the ability of Pavlov's dogs to salivate when the bells rang-seems to be located in the limbic system. This is long-term memory, but of a very restricted kind. The sophisticated sort of human long-term memory is situated in the neocortex, which is consistent with the human ability to think ahead. As we grow old, we sometimes forget what has just been said to us while retaining vivid and accurate recollections of events in our childhood. In such cases, little seems to be wrong with either our short-term or our long-term memories; the problem is the connection between the two-we have great difficulty in accessing new material into the long-term memory. Penfield believed that this lost accessing ability arises from an inadequate blood supply to the hippocampus in old age-because of arteriosclerosis or other physical disabilities. Thus elderly people-and ones not so elderly-may have serious impairments in accessing short-term memory while being otherwise perfectly alert and intellectually keen.* This phenomenon also shows a clear-cut distinction between short-term and long-term memory, consistent with their localization in different parts of the brain. Waitresses in short-order restaurants can remember an impressive amount of information, which they accurately transmit to the kitchen. But an hour later, the information has vanished utterly. It was put into the short-term memory only, and no effort was made to access it into the long-term memory.
The mechanics of recall can be complex. A common experience is that we know something is in our long-term memory-a word, a name, a face, an experience-but find ourselves unable to call it up. No matter how hard we try, the memory resists retrieval. But if we think sideways at it, recalling some slightly related or peripheral item, it often follows unbidden. (Human vision is also a little like this. When we look directly at a faint object-a star, say-we are using the fovea, the part of the retina with the greatest acuity and the greatest density of cells called cones. But when we avert our vision slightly-when, in a manner of speaking, we look sideways at the object-we bring into play the cells called rods, which are sensitive to feeble illumination and so able to perceive the faint star.) It would be interesting to know why thinking sideways improves memory retrieval; it may be merely associating to the memory trace by a different neural pathway. But it does not suggest particularly efficient brain engineering.
* Indeed, there is a range of medical evidence on the connection between blood supply and intellectual abilities. It has long been known that patients deprived of oxygen for some minutes can experience permanent and serious mental impairment. Operations to remove material from clogged carotid arteries in an effort to prevent stroke yield unexpected benefits. According to one study, six weeks after such operations, the patients showed an average increase in IQ of eighteen points, a substantial improvement. And there has been some speculation that immersion in hyperbaric oxygen-that is, oxygen under high pressure-can raise the intelligence of infants.
We have all had the experience of awakening with a particularly vivid, chilling, insightful or otherwise memorable dream clearly in mind; saying to ourselves, "I'll certainly remember this dream in the morning"; and the next day having not the foggiest notion about the content of the dream or, at best, a vague trace of an emotion tone. On the other hand, if I am sufficiently exercised about the dream to awaken my wife in the middle of the night and tell her about it, I have no difficulty remembering its contents unaided in the morning. Likewise, if I take the trouble of writing the dream down, when I awaken the next morning I can remember the dream perfectly well without referring to my notes. The same thing is true of, for example, remembering a telephone number. If I am told a number and merely think about it, I am likely to forget it or transpose some of the digits. If I repeat the numbers out loud or write them down, I can remember them quite well. This surely means that there is a part of our brain which remembers sounds and images, but not thoughts. I wonder if that sort of memory arose before we had very many thoughts-when it was important to remember the hiss of an attacking reptile or the shadow of a plummeting hawk, but not our own occasional philosophical reflections.
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