Human brain

This article is about features specific to the human brain. For basic information about brains, see Brain.

Human brain
Human brain and skull
Cerebral lobes: the frontal lobe (pink), parietal lobe (green) and occipital lobe (blue)
Details
Precursor	Neural tube
System	Central nervous system Neuroimmune system
Artery	Internal carotid arteries, vertebral arteries
Vein	Internal jugular vein, internal cerebral veins, external veins: (superior and inferior cerebral veins, and middle cerebral veins), basal vein, terminal vein, choroid vein, cerebellar veins
Identifiers
Latin	Cerebrum[1]
Greek	ἐγκέφαλος (enképhalos)[2]
MeSH	D001921
TA	14.1.03.001
FMA	50801
Anatomical terminology [edit on Wikidata]

The human brain is the main organ of the human central nervous system. It is located in the head, protected by the skull. It has the same general structure as the brains of other mammals, but with a more developed cerebral cortex. Large animals such as whales and elephants have larger brains in absolute terms, but when measured using a measure of relative brain size, which compensates for body size, the quotient for the human brain is almost twice as large as that of a bottlenose dolphin, and three times as large as that of a chimpanzee, though the quotient for a treeshrew's brain is larger than that of a human's.^[3] Much of the size of the human brain comes from the cerebral cortex, especially the frontal lobes, which are associated with executive functions such as self-control, planning, reasoning, and abstract thought. The area of the cerebral cortex devoted to vision, the visual cortex, is also greatly enlarged in humans compared to other animals.

The human cerebral cortex is a thick layer of neural tissue that covers the two cerebral hemispheres that make up most of the brain. This layer is folded in a way that increases the amount of surface area that can fit into the volume available. The pattern of folds is similar across individuals but shows many small variations. The cortex is divided into four lobes – the frontal lobe, parietal lobe, temporal lobe, and occipital lobe. (Some classification systems also include a limbic lobe and treat the insular cortex as a lobe.) Within each lobe are numerous cortical areas, each associated with a particular function, including vision, motor control, and language. The left and right hemispheres are broadly similar in shape, and most cortical areas are replicated on both sides. Some areas, though, show strong lateralization, particularly areas that are involved in language. In most people, the left hemisphere is dominant for language, with the right hemisphere playing only a minor role. There are other functions, such as visual-spatial ability, for which the right hemisphere is usually dominant.

Despite being protected by the thick bones of the skull, suspended in cerebrospinal fluid, and isolated from the bloodstream by the blood–brain barrier, the human brain is susceptible to damage and disease. The most common forms of physical damage are closed head injuries such as a blow to the head or other trauma, a stroke, or poisoning by a number of chemicals that can act as neurotoxins, such as alcohol. Infection of the brain, though serious, is rare because of the protective blood-to brain and blood-to cerebral fluid barriers. The human brain is also susceptible to degenerative disorders, such as Parkinson's disease, forms of dementia including Alzheimer's disease, (mostly as the result of aging) and multiple sclerosis. A number of psychiatric conditions, such as schizophrenia and clinical depression, are thought to be associated with brain dysfunctions, although the nature of these is not well understood. The brain can also be the site of brain tumors and these can be benign or malignant.

There are some techniques for studying the brain that are used in other animals that are not suitable for use in humans and vice versa; it is easier to obtain individual brain cells taken from other animals, for study. It is also possible to use invasive techniques in other animals such as inserting electrodes into the brain or disabling certains parts of the brain in order to examine the effects on behaviour – techniques that are not possible to be used in humans. However, only humans can respond to complex verbal instructions or be of use in the study of important brain functions such as language and other complex cognitive tasks, but studies from humans and from other animals, can be of mutual help. Medical imaging technologies such as functional neuroimaging and EEG recordings are important techniques in studying the brain. The complete functional understanding of the human brain is an ongoing challenge for neuroscience.

 

Contents

Structure

General features

 

The adult human brain weighs on average about 1.3–1.5 kg (2.9–3.3 lb), or about 2% of total body weight,^[4][5] with a volume of around 1130 cm³ in women and 1260 cm³ in men, although there is substantial individual variation.^[6] Neurological differences between the sexes have not been shown to correlate in any simple way with IQ or other measures of cognitive performance.^[7]

The human brain is composed of neurons, glial cells, and blood vessels. The number of neurons is estimated at roughly 100 billion.^[8] The adult human brain is estimated to contain 86±8 billion neurons, with a roughly equal number (85±10 billion) of non-neuronal cells. Out of these, 16 billion (or 19% of all brain neurons) are located in the cerebral cortex (including subcortical white matter), 69 billion (or 80% of all brain neurons) are in the cerebellum.^[5][9]

The cerebral hemispheres (the cerebrum) form the largest part of the human brain and are situated above other brain structures. They are covered with a cortical layer (the cerebral cortex) which has a convoluted topography.^[10] Underneath the cerebrum lies the brainstem, resembling a stalk on which the cerebrum is attached. At the rear of the brain, beneath the cerebrum and behind the brainstem, is the cerebellum, a structure with a horizontally furrowed surface, the cerebellar cortex, that makes it look different from any other brain area. The same structures are present in other mammals, although they vary considerably in relative size. As a rule, the smaller the cerebrum, the less convoluted the cortex. The cortex of a rat or mouse is almost perfectly smooth. The cortex of a dolphin or whale, on the other hand, is more convoluted than the cortex of a human.

The living brain is very soft, having a gel-like consistency similar to soft tofu. Although referred to as grey matter, the live cortex is pinkish-beige in color and slightly off-white in the interior.

Comparative anatomy

 

The human brain has many properties that are common to all vertebrate brains, including a basic division into three parts called the forebrain, midbrain, and hindbrain, with interconnected fluid-filled ventricles, and a set of generic vertebrate brain structures including the medulla oblongata and pons of the brainstem, the cerebellum, optic tectum, thalamus, hypothalamus, basal ganglia, olfactory bulb, and many others.

As a mammalian brain, the human brain has special features that are common to all mammalian brains, most notably a six-layered cerebral cortex and a set of associated structures, including the hippocampus and amygdala. The upper surface of the forebrain of other vertebrates is covered in a layer of neural tissue called the pallium. The pallium is a relatively simple three-layered cell structure. The hippocampus and the amygdala originate from the pallium but in mammals they are much more complex.

As a primate brain, the human brain has a much larger cerebral cortex, in proportion to body size, than most mammals, and a very highly developed visual system. The shape of the brain within the skull is also altered somewhat as a consequence of the upright position in which primates hold their heads.

As a hominid brain, the human brain is substantially enlarged even in comparison to the brain of a typical monkey. The sequence of evolution from Australopithecus (four million years ago) to Homo sapiens (modern man) was marked by a steady increase in brain size, particularly in the frontal lobes, which are associated with a variety of high-level cognitive functions.

Humans and other primates have some differences in gene sequence, and genes are differentially expressed in many brain regions. The functional differences between the human brain and the brains of other animals also arise from many gene–environment interactions.^[11]

The neuroimmune system of the brain is structurally distinct from the peripheral immune system which protects the rest of the body; in particular, the immune system is composed primarily of hematopoietic cells and anatomical barriers, while the neuroimmune system is composed of glia, mast cells, and various brain barriers (e.g., blood–brain barrier and blood-cerebrospinal fluid barrier).

Cerebral cortex

 

The dominant feature of the human brain is corticalization. The cerebral cortex in humans is so large that it overshadows every other part of the brain. A few subcortical structures show alterations reflecting this trend. The cerebellum, for example, has a medial zone connected mainly to subcortical motor areas, and a lateral zone connected primarily to the cortex. In humans the lateral zone takes up a much larger fraction of the cerebellum than in most other mammalian species. Corticalization is reflected in function as well as structure. In a rat, surgical removal of the entire cerebral cortex leaves an animal that is still capable of walking around and interacting with the environment.^[13]

In a human, comparable cerebral cortex damage produces a permanent state of coma. The amount of association cortex, relative to the other two categories of sensory and motor, increases dramatically as one goes from simpler mammals, such as the rat and the cat, to more complex ones, such as the chimpanzee and the human.^[14] A gene present in the human genome but not in the chimpanzee (ArhGAP11B) seems to play a major role in corticalization. ArhGAP11B and human encephalisation The cerebral cortex is essentially a sheet of neural tissue, folded in a way that allows a large surface area to fit within the confines of the skull. When unfolded, each cerebral hemisphere has a total surface area of about 1.3 square feet (0.12 m²).^[15] Each cortical ridge is called a gyrus, and each groove or fissure separating one gyrus from another is called a sulcus.

Cortical divisions

 

The cerebral cortex is nearly symmetrical with left and right hemispheres that are approximate mirror images of each other. Each hemisphere is conventionally divided into four "lobes", the frontal lobe, parietal lobe, occipital lobe, and temporal lobe. With one exception, this division into lobes does not derive from the structure of the cortex, though the lobes are named after the bones of the skull that overlie them, the frontal bone, parietal bone, temporal bone, and occipital bone. The borders between lobes lie beneath the sutures that link the skull bones together. The exception is the border between the frontal and parietal lobes, which lies behind the corresponding suture; instead it follows the anatomical boundary of the central sulcus, a deep fold in the brain's structure where the primary somatosensory cortex and primary motor cortex meet.

Because of the arbitrary way most of the borders between lobes are demarcated, they have little functional significance. With the exception of the occipital lobe, a small area that is entirely dedicated to vision, each of the lobes contains a variety of brain areas that have minimal functional relationship. The parietal lobe, for example, contains areas involved in somatosensation, hearing, language, attention, and spatial cognition. In spite of this heterogeneity, the division into lobes is convenient for reference. The main functions of the frontal lobe are to control attention, abstract thinking, behavior, problem solving tasks, and physical reactions and personality. The occipital lobe is the smallest lobe; its main functions are visual reception, visual-spatial processing, movement, and color recognition. The temporal lobe controls auditory and visual memories, language, and some hearing and speech.

 

 

 

Although there are enough variations in the shape and placement of gyri and sulci (cortical folds) to make every brain unique, most human brains show sufficiently consistent patterns of folding that allow them to be named. Many of the gyri and sulci are named according to the location on the lobes or other major folds on the cortex. These include:

• Superior, Middle, Inferior frontal gyrus: in reference to the frontal lobe
• Medial longitudinal fissure, which separates the left and right cerebral hemispheres
• Precentral and Postcentral sulcus: in reference to the central sulcus, which separates the frontal lobe from the parietal lobe
• Lateral sulcus, which divides the frontal lobe and parietal lobe above from the temporal lobe below
• Parieto-occipital sulcus, which separates the parietal lobes from the occipital lobes, is seen to some small extent on the lateral surface of the hemisphere, but mainly on the medial surface.
• Trans-occipital sulcus: in reference to the occipital lobe

Functional divisions

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Functions of the cortex are divided it into three categories of regions: One consists of the primary sensory areas, which receive signals from the sensory nerves and tracts by way of relay nuclei in the thalamus. Primary sensory areas include the visual area of the occipital lobe, the auditory area in parts of the temporal lobe and insular cortex, and the somatosensory cortex in the parietal lobe. A second category is the primary motor cortex, which sends axons down to motor neurons in the brainstem and spinal cord. This area occupies the rear portion of the frontal lobe, directly in front of the somatosensory area. The third category consists of the remaining parts of the cortex, which are called the association areas. These areas receive input from the sensory areas and lower parts of the brain and are involved in the complex processes of perception, thought, and decision-making.^[16]

Cytoarchitecture

 

Different parts of the cerebral cortex are involved in different cognitive and behavioral functions. The differences show up in a number of ways: the effects of localized brain damage, regional activity patterns exposed when the brain is examined using functional imaging techniques, connectivity with subcortical areas, and regional differences in the cellular architecture of the cortex. Neuroscientists describe most of the cortex—the part they call the neocortex—as having six layers, but not all layers are apparent in all areas, and even when a layer is present, its thickness and cellular organization may vary. Scientists have constructed maps of cortical areas on the basis of variations in the appearance of the layers as seen with a microscope. One of the most widely used schemes came from Korbinian Brodmann, who split the cortex into 51 different areas and assigned each a number (many of these Brodmann areas have since been subdivided). For example, Brodmann area 1 is the primary somatosensory cortex, Brodmann area 17 is the primary visual cortex, and Brodmann area 25 is the anterior cingulate cortex.^[17]

Topography

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Many of the brain areas Brodmann defined have their own complex internal structures. In a number of cases, brain areas are organized into "topographic maps", where adjoining bits of the cortex correspond to adjoining parts of the body, or of some more abstract entity. A simple example of this type of correspondence is the primary motor cortex, a strip of tissue running along the anterior edge of the central sulcus, shown in the image to the right. Motor areas innervating each part of the body arise from a distinct zone, with neighboring body parts represented by neighboring zones. Electrical stimulation of the cortex at any point causes a muscle-contraction in the represented body part. This "somatotopic" representation is not evenly distributed, however. The head, for example, is represented by a region about three times as large as the zone for the entire back and trunk. The size of any zone correlates to the precision of motor control and sensory discrimination possible. The areas for the lips, fingers, and tongue are particularly large, considering the proportional size of their represented body parts.

In visual areas, the maps are retinotopic—that is, they reflect the topography of the retina, the layer of light-activated neurons lining the back of the eye. In this case too the representation is uneven: the fovea—the area at the center of the visual field—is greatly overrepresented compared to the periphery. 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. 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. Visual areas farther downstream extract features such as color, motion, and shape.

In auditory areas, the primary map is tonotopic. Sounds are parsed according to frequency (i.e., high pitch vs. low pitch) by subcortical auditory areas, and this parsing is reflected by the primary auditory zone of the cortex. As with the visual system, there are a number of tonotopic cortical maps, each devoted to analyzing sound in a particular way.

Within a topographic map there can sometimes be finer levels of spatial structure. In the primary visual cortex, for example, where the main organization is retinotopic and the main responses are to moving edges, cells that respond to different edge-orientations are spatially segregated from one another.

Development

During the first three weeks of gestation, the human embryo's ectoderm forms a thickened strip called the neural plate. The neural plate then folds and closes to form the neural tube. This tube flexes as it grows, forming the crescent-shaped cerebral hemispheres at the head, and the cerebellum and pons towards the tail.

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