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The brain's cerebral cortex, or neocortex, has specialized areas for language, sensory, and emotion processing.
In Evolving Brains (2000), John Allman explains that cortex means "the outer shell or rind of an object." He explains that the prefix "neo" implies that it is new. Allman writes: "The neocortex is found only in mammals, although it is related to forebrain structures found in other vertebrate classes. The neocortex, the sheetlike, six-layered structure in the roof of the forebrain that is found in all mammals and only in mammals, was probably present in the earliest true mammals; it is possible that it may actually have evolved earlier, at some point after the separation of the line leading to the mammals from the lines leading to reptiles and birds. Jaak Panksepp, in Affective Neuroscience: The Foundations of Human and Animal Emotions (1998), explains that "the modular unit of the neocortex is a column, a vertically oriented functional grouping of about 4,000 interconnected neurons with comparatively weak connections to immediately neighboring modules.
Before I discuss the anatomical differentiations referred to as lobes, I would like to discuss a most important recessed area of cortex that traditionally has been considered as part of the limbic system or paleomammalian brain.
The cingulate cortex shapes itself to the outside curve of the corpus callosum, the thick band of nerve fibers that connects the brain's left and right hemispheres (see The Brain's Two Hemispheres).
Damasio calls attention to an interesting phenomenon that illustrates the function of the anterior cingulate cortex. To clarify where emotional expression does originate, Damasio writes: "If you study a patient in whom a stroke has damaged the anterior cingulate in the left hemisphere you will see precisely the opposite result. Paul MacLean made a very interesting observation relating to the cingulate cortex and the bond between mother and child in an article included in the book The Limbic System and Clinical Disorders (B.K.
The anterior cingulate cortex may play an important role in bonds between parent and child, as well as other attachments. Wikipedia's entry for the "Human brain" states: "the lobes are named after the bones of the skull that overlie them.
The occipital lobe includes sophisticated topographical maps with complex interconnections required for visual processing, and according to MedlinePlus Dictionary, has "the form of a 3-sided pyramid." The Merck Manuals Online Medical Library states that in addition to processing and interpreting vision, cortical areas in the occipital lobe enable people to form visual memories and integrate visual perceptions with spatial information coming from the adjacent parietal lobes. The parietal lobe contains an area that processes bodily sensations—the somatosensory cortex, which we will discuss further later in this narrative.
A fifth lobe, the insula, is located "in the center of the cerebral hemisphere that is situated deeply between the lips of the sylvian fissure," according to MedlinePlus Dictionary.
The illustration to the left (image links to source) is a pyramidal cell, which is common in the prefrontal cortex. In his video course, Sapolsky explains that the frontal cortex "helps you focus on what the task is right now.
Temple Grandin and Catherine Johnson make a good point in Animals in Translation: Using the Mysteries of Autism to Decode Animal Behavior (2005). The importance of the brain—especially frontal regions of the neocortex—in shaping behavior and personality began to be realized in the aftermath of an astonishing injury to a man named Phineas Gage in 1848, then a reliable and successful construction foreman working on the railroad in Vermont.
Macmillan provides all the unpleasant but extremely interesting details of how physician John Martyn Harlow treated Gage's injury. Henry Jacob Bigelow, who presented Gage and his tamping iron at an 1849 meeting of the Boston Society of Medical Improvement. According to Harlow, Gage was tamping the powder and fuse ("slightly" in the 1848 account) prior to the sand's being poured in when his attention was attracted by his men, who were loading excavated rock onto a platform car in the pit a few feet behind him. During Gage's recovery period, Macmillan details how Harlow began to notice unusual behavior. Macmillan explains that what we know of Gage's employment subsequent to the accident comes from Harlow. The gold rush and its associated influx of people into California, explains Macmillan, may be responsible for what Gage did in 1852. In detailing how the Gage case shaped thinking about localization of brain function, Macmillan calls attention to the work of David Ferrier (1843-1928). The mind exercises a constant inhibitory influence upon all action, physical or mental; from the simple restraint upon the lower reflexes, such as the action of the sphincters, to the higher control over the complex reflexes, such as emotional impulses and their manifestation in speech and expression.
In the twenty-first century, we better understand that the role of specialized areas of the neocortex—the frontal lobes especially including the orbital-frontal cortex—is to modulate or "inhibit" emotion-driven behavior originating in subcortical mammalian brain structures. In most humans, especially right-handed humans, specialized cortical language areas are located in the brain's left hemisphere.
The Canadian Institute of Neurosciences, Mental Health and Addiction, and the Canadian Institutes of Health Research, provides the image to the left, which links to an educational module, The Brain from Top to Bottom. Subsequent brain imaging experiments have revealed a third region of the brain—the inferior parietal lobule—that large bundles of nerve fibers connect to both Broca's area and Wernicke's area.
The image to the left is also taken from the intermediate-level module of The Brain from Top to Bottom (image links to source).
Damage to the right hemisphere illustrates its vital role in communication as well as the complex connectedness of the brain.
In anosognosia, which Damasio discusses in Descartes' Error, an individual cannot recognize certain parts of their body as being their own. Right-hemisphere damage also can affect an individual's use of prosody, which is the ability to use intonation and stress to convey the emotions one feels. In Brainscapes, Restak illuminates what happens when the language-processing left hemisphere is taken out of the loop. In a module taken from The Brain from Top to Bottom, you can find the following excellent summary of Brodmann's Areas: "The cellular architecture differs sufficiently from one part of the neocortex to another to be used as a criterion for defining cortical areas that are functionally distinct. The Canadian internet resource referenced above went on to explain: "Brodmann's intuition, whose accuracy has been confirmed many times since, was that a particular anatomical structure corresponded to a particular function. In the Brodmann's Areas illustration to the left, the primary motor cortex corresponds to the area labeled 4. The Brain from Top to Bottom, the areas of cortex assigned to various body parts "are proportional not to their size, but rather to the complexity of the movements that they can perform. In Evolving Brains, Allman cites the work of Hughlings Jackson in determining how the brain controls muscle movement. Allman explains that Jackson's observations "relate to three fundamental properties of the neocortex.
In the illustration to the left (image links to source), areas labeled 3, 1, and 2 represent the somatosensory cortex, another kind of topographical map. Regarding the role of experience in determining cortical representations, Allman writes: "Functional imaging experiments done in human subjects have also demonstrated that the hand representation expands as a result of performing complex finger movements.
Although Allman eloquently covers visual processing in Evolving Brains, at this point in my research, the material is a bit over my head, to say the least.
In the Russo-Japanese War of 1905, many Japanese soldiers sustained bullet wounds that penetrated through the posterior part of their brains.
In The Modular Brain: How New Discoveries in Neuroscience are Answering Age-Old Questions about Memory, Free Will, Consciousness, and Personal Identity (1994), Richard M. As a last note on the vastly complex visual cortical areas, in Evolving Brains, Allman notes the differences in the visual cortex that distinguish earlier mammals from primates and humans. The antecedents of the neocortex are present in the telencephalic roof in even the most primitive vertebrates.


These columns are strongly linked to other cortical modules and to lower brain areas by descending and looping connections. I am trying to avoid specifying specific components of what has in the past been called the limbic system since a discrete system does not exist. MedlinePlus Dictionary defines anterior as "relating to or situated near or toward the head… ." Thus, the anterior cingulate cortex in the image above, outlined in red, bumps up against the inside of our frontal lobe. Damage to the motor cortex will affect a voluntary smile but not a genuine, emotional smile generated from within the anterior cingulate cortex. In repose or in emotion-related movement, the face is asymmetrical, less mobile on the right than on the left. Nestled within each temporal lobe are the amygdala and the hippocampus, which are, writes Restak in Brainscapes, "involved in learning, memory and the experience and expression of emotion." A portion of the temporal lobe, called the entorhinal cortex, channels cortical inputs to the hippocampus. To get an idea of where the insula is located, the image to the right (links to source) illustrates the location of the sylvian fissure. This lobe plays a role in certain language functions and when damaged can lead to aphasia, the inability to use or understand spoken and written language. Bob Jacobs, Laboratory of Quantitative Neuromorphology, Department of Psychology, Colorado College, provides this photograph. Early state dementia patients asked to count backward from 20 will start the task and then revert to reciting the months of the year.
In the next subsection, we will discuss a particular portion of the frontal cortex, the orbital-frontal cortex, so labeled in the illustration to the right (links to source). If you damage any part of your brain in an accident or a stroke, they explain, it may appear that you have damage to your frontal lobes, even when your frontal lobes remain perfectly intact.
The location of this most important region is highlighted in the image above right (links to source; this image is an MRI of Wikipedia contributor Paul Wicks's brain). Gage was using a tamping iron to compact material—including explosive powder that at some point in time is topped with sand—into a hole drilled into a bed of rock when he accidentally set off an explosion that sent a tamping iron through his head. With his head still averted and while continuing to look over his right shoulder, Gage dropped the iron onto the powder again where, this time, it hit the rock, struck a spark, ignited the charge, and immediately reversed its initial direction.
Regarding his patient, Harlow wrote: "Does not estimate size or money accurately, though he has memory as perfect as ever.
After spending some time displaying his injury and tamping iron in a museum setting, Harlow wrote that Gage began working with horses.
Allen Starr published "the first modern comparison of the effects of different kinds of [brain] lesions and tumor" and that Gage figured prominently in the report.
This action of control implies a recognition of the import of an act in connection with other acts; in a word, it involves judgment and reason, the highest mental qualities. In many left-handed people, however, specialized cortical language areas exist in both hemispheres. The particular module to which this image links provides a good overview of Paul Broca and Carl Wernicke, two nineteenth-century neuroanatomists who, during autopsies, studied the brains of people who had suffered from language disorders during their lives.
This module explains that the left hemisphere controls the "phonological, syntactic, and lexical aspects" of conversation. In fact, these people react exactly as if they were dealing with idioms in a foreign language: their grammar and phonology may be correct, but they do not understand the verbal humour or metaphors that native speakers of that language use every day. The image to the right—taken while the subject was generating words—links to an advanced-level module of The Brain from Top to Bottom. That is what the German anatomist Korbinian Brodmann did in the early 20th century, when he developed a map of the brain based on the differences in the cellular architecture of the various parts of the cortex. For example, Brodmann's area 17, which receives information from a nucleus of the thalamus that is connected to the retina, turns out to correspond precisely to the primary visual cortex. The neurosurgeon Wilder Graves Penfield (1891-1976) explored this region while treating patients with severe epilepsy in Montreal. Hence, the areas for the hand and face are especially large compared with those for the rest of the body.
The homunculus to the right (image links to source) displays the proportion of cortex dedicated to controlling different parts of the body. If your dog licks the bottom of your foot, tickling it, a specific area of the cortex will be activated. The expansion of the hand representation can be observed following short-term training, but it is most notable in Braille readers and in musicians who play stringed instruments. The visual circuitry in the human cerebral cortex contains several dozen distinct retinotopic maps, each devoted to analyzing the visual input stream in a particular way.
For orientation as to where these visual areas area, please reference the illustration to the left. Because of the higher muzzle velocity and the smaller bullet size of rifles developed in the late nineteenth century, these weapons tended to produce more localized brain injuries than were inflicted in earlier wars, and improved care of the wounded also resulted in higher rates of survival. Restak provides an example of how damage to one area of the brain, in only one hemisphere, can dramatically change how we perceive the world. He writes that "opossums and hedgehogs, which in many respects resemble the early mammals that lived more than 60 million years ago, have rather limited visual capacities and a small number of the visual cortical areas. The neocortex is a specialization in the telencephalon that parallels the formation of the dorsal ventricular ridge and wulst in reptiles and birds. I think the point to remember is that the anterior cingulate cortex is more ancient, from an evolutionary perspective, than outer areas of cortex. But if the patient tries to contract the facial muscles willfully, the movements are carried out normally and symmetry returns. If you jump ahead to this link, remember to click the BACK button on your browser to return here. While we will discuss specialized areas of the neocortex later in this narrative, we will discuss the lobes in brief below.
In reading, however, you are likely to come across other terminology that designates areas of the brain slightly different from that pictured above. By inhibiting all but one set of impulses it enables one to fix attention upon a subject, and hold it there. Under optimal circumstances, our large human fontal lobes inspire us to adjust behavior based on environmental circumstances as well as familial, cultural, and societal influences.
A minority of both left- and right-handed people appear to have specialized cortical areas for language in only the right hemisphere.
Broca and Wernicke were thus able to correlate damage to specific areas with language deficits. The image to the right links to an intermediate-level module in The Brain from Top to Bottom that sums up the important role of this area: "The inferior parietal lobule of the left hemisphere lies at a key location in the brain, at the junction of the auditory, visual, and somatosensory cortexes, with which it is massively connected. This explains why the left hemisphere was long considered the dominant hemisphere for language. This module explains that right-hemisphere damage can result in hemineglect, in which one pays no attention to stimuli coming in from the left side of the body.


This mapping allows you to know that it is your foot being licked, not the back of your neck. Meanwhile, I will provide here some basic information about cortical maps involved in visual processing. The primary visual cortex (Brodmann area 17), which is the main recipient of direct input from the visual part of the thalamus, contains many neurons that are most easily activated by edges with a particular orientation moving across a particular point in the visual field.
Many of the wounded soldiers were partially blinded by these injuries, and Tatsuji Inouye, an ophthalmologist, was asked by the Japanese government to evaluate the extent of their blindness as a means to determine their pension benefits.
Restak discusses how Michael Gazzaniga conducted an experiment that involved a subject who had lost sight off to his left because of brain damage to the visual area on the right side of his brain. In these mammals, the cortical maps of the retina are relatively uniform in that the amount of cortical space devoted to the more central part of the visual field in front of the animal is not much greater than the cortex devoted to the more peripheral parts of the visual field.
Jones and other scientists from the Center for Neuroscience, University of California, Davis and University of Wisconsin, Madison.
The neocortex is just as much a unique defining feature of mammals as are the mammary glands or the malleus and incus in the middle ear. But a neuropsychologist named Elkhonon Goldberg at New York University School of Medicine, who wrote a fantastic book about frontal lobe functions called The Executive Brain, has a different theory. Damasio, for example, clarifies his use of terms in Descartes' Error: "In neuroanatomical terminology, the orbital region is known also as the ventromedial region of the frontal lobe, and this is how I [Damasio] will refer to it throughout the book. The image links to their web site, Meet Phineas Gage.) Although Gage was not able to return to work as a foreman, he did pursue and obtain other employment relating to the care and management of horses. According to Bigelow, the powder and fuse had already been "adjusted" in the hole, and Gage had instructed an assistant to pour in the sand. Although we can survive without certain components of the neocortex, we are subsequently very different in terms of personality, temperament, and judgement. Wikipedia aptly describes the brain's plasticity: "Studies of children have shown that if a child has damage to the left hemisphere, the child may develop language in the right hemisphere instead. In addition, the neurons in this lobule have the particularity of being multimodal, which means that they can process different kinds of stimuli (auditory, visual, sensorimotor, etc.) simultaneously.
The left hemisphere is not, however, solely responsible for communication and the module goes on to explain the role of the right hemisphere. While the patient was under local anesthesia, Penfield stimulated the brain with electrical probes and observed the patient's responses. Brodmann's area 18 represents the secondary visual cortex (V2) and Brodmann's area 19 represents the associative visual cortex (V3). Inouye found that the part of the visual field in which these soldiers were blind corresponded to the locations of their brain injuries as determined by the sites of the bullet's entry and exit through the head.
The subject "was asked to imagine himself looking toward California from New York and naming the states in between. By contrast, primates have extremely well developed visual capacities and have a large number of cortical maps devoted to visual perception and memory. As with the other distinctive features of mammals, the neocortex probably evolved as a part of a set of adaptations related to temperature homeostasis. Anatomists call each cortical fold a sulcus and the smooth protruding area between folds a gyrus.
Yet when the patient smiles or laughs spontaneously, in response to a humorous remark, something entirely different happens: the smile is normal, both sides of the face move as they should, and the expression is natural, no different from the usual pre-paralysis smile of that individual.
He thinks that while the frontal lobes may be more fragile, there is another factor involved, which is that every other part of the brain is connected to them. After learning something about brain evolution and ethology, I am not surprised that in this subsequent work Gage engaged with animals.
While waiting for this to be done, Gage turned his head away, and after an interval of a few seconds again dropped the iron, this time, as he supposed, onto the sand.
This combination of traits makes the inferior parietal lobule an ideal candidate for apprehending the multiple properties of spoken and written words: their sound, their appearance, their function, etc. He did this to identify areas requiring surgery and to avoid vital areas that should not be destroyed.
By combining the visual field deficits from different soldiers he was able to deduce the topographic organization of the primary visual cortex.
The large increases in metabolic expenditure necessary to sustain temperature homeostasis required commensurate increases in the acquisition of food by the early mammals.
This illustrates that the motor control for an emotion-related movement sequence is not in the same location as the control for a voluntary act.
I will again quote Damasio to forestall the impulse to credit this or that lobe with a discrete process. When you damage any part of the brain, you change input to the frontal lobes, and when you change input, you change output.
But before we get to this topic, lets review a few facts about the accident and Gage's medical treatment, taken from the well researched An Odd Kind of Fame: Stories of Phineas Gage (2002), by Malcolm Macmillan. However, no sand had been added, and, when the bar struck the rock, the charge exploded, driving it upward. The right hemisphere provides the ability to go beyond the literal meanings of words and employs multiple processes to do so. In doing this, he observed that stimulation to certain areas of the cortex triggered highly localized muscle contractions on the opposite side of the body. Inouye's map revealed that much more cortex was devoted to the representation of the central part of the retina than to the periphery. He omitted the states to the left, corresponding to the visual field mediated by his right brain lesion. Since these animals were small and had only a limited capacity to store energy as fat, they were constantly under the threat of starvation.
If the frontal lobes aren't getting the right input, they don't produce the right output even though structurally they're fine. As mentioned earlier when discussing the temporal lobe, Brodmann's areas 20 and 21 in the temporal lobe (also circled in red) represent the inferotemporal visual cortex, which includes neurons sensitive to the images of faces and sends visual information to the amygdala. This is the portion of the retina with the highest acuity, and it is our most important means for probing our environment for information, and the part you are using to read this book.
The neocortex stores information about the structure of the environment so that the mammal can readily find food and other resources necessary for its survival (emphasis added).
Inouye's map of the primary visual cortex has been confirmed by modern brain-imaging techniques.



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