In human neuroanatomy, brain asymmetry can refer to at least two quite distinct findings:
Neuroanatomical differences themselves exist on different scales, from neuronal densities, to the size of regions such as the planum temporale, to—at the largest scale—the torsion or "wind" in the human brain, reflected shape of the skull, which reflects a backward (posterior) protrusion of the left occipital bone and a forward (anterior) protrusion of the right frontal bone. [2] In addition to gross size differences, both neurochemical and structural differences have been found between the hemispheres. Asymmetries appear in the spacing of cortical columns, as well as dendritic structure and complexity. Larger cell sizes are also found in layer III of Broca's area.
The human brain has an overall leftward posterior and rightward anterior asymmetry (or brain torque). There are particularly large asymmetries in the frontal, temporal and occipital lobes, which increase in asymmetry in the antero-posterior direction beginning at the central region. Leftward asymmetry can be seen in the Heschl gyrus, parietal operculum, Silvian fissure, left cingulate gyrus, temporo-parietal region and planum temporale. Rightward asymmetry can be seen in the right central sulcus (potentially suggesting increased connectivity between motor and somatosensory cortices in the left side of the brain), lateral ventricle, entorhinal cortex, amygdala and temporo-parieto-occipital area. Sex-dependent brain asymmetries are also common. For example, human male brains are more asymmetrically lateralized than those of females. However, gene expression studies done by Hawrylycz and colleagues and Pletikos and colleagues, were not able to detect asymmetry between the hemispheres on the population level. [3] [4]
People with autism have much more symmetrical brains than people without it. [5] [6]
In the mid-19th century scientists first began to make discoveries regarding lateralization of the brain, or differences in anatomy and corresponding function between the brain's two hemispheres. Franz Gall, a German anatomist, was the first to describe what is now known as the Doctrine of Cerebral Localization. Gall believed that, rather than the brain operating as a single, whole entity, different mental functions could be attributed to different parts of the brain. He was also the first to suggest language processing happened in the frontal lobes. [7] However, Gall's theories were controversial among many scientists at the time. Others were convinced by experiments such as those conducted by Marie-Jean-Pierre Flourens, in which he demonstrated lesions to bird brains caused irreparable damage to vital functions. [8] Flourens's methods, however, were not precise; the crude methodology employed in his experiments actually caused damage to several areas of the tiny brains of the avian models.
In 1861, surgeon Paul Broca provided evidence that supported Gall's theories. Broca discovered that two of his patients who had had speech loss had similar lesions in the same area of the left frontal lobe. [7] While this was compelling evidence for localization of function, the connection to "sidedness" was not made immediately. As Broca continued to study similar patients, he made the connection that all of the cases involved damage to the left hemisphere, and in 1864 noted the significance of these findings—that this must be a specialized region. He also—incorrectly—proposed theories about the relationship of speech areas to "handedness".
Accordingly, some of the most famous early studies on brain asymmetry involved speech processing. Asymmetry in the Sylvian fissure (also known as the lateral sulcus), which separates the frontal and parietal lobes from the temporal lobe, was one of the first incongruencies to be discovered. Its anatomical variances are related to the size and location of two areas of the human brain that are important for language processing, Broca's area and Wernicke's area, both in the left hemisphere. [9]
Around the same time that Broca and Wernicke made their discoveries, neurologist Hughlings Jackson suggested the idea of a "leading hemisphere"—or, one side of the brain that played a more significant role in overall function—which would eventually pave the way for understanding hemispheric "dominance" for various processes. Several years later, in the mid-20th century, critical understanding of hemispheric lateralization for visuospatial, attention and perception, auditory, linguistic and emotional processing came from patients who underwent split-brain procedures to treat disorders such as epilepsy. In split-brain patients, the corpus callosum is cut, severing the main structure for communication between the two hemispheres. The first modern split-brain patient was a war veteran known as Patient W.J., [10] whose case contributed to further understanding of asymmetry.
Brain asymmetry is not unique to humans. In addition to studies on human patients with various diseases of the brain, much of what is understood today about asymmetries and lateralization of function has been learned through both invertebrate and vertebrate animal models, including zebrafish, pigeons, rats, and many others. For example, more recent studies revealing sexual dimorphism in brain asymmetries in the cerebral cortex and hypothalamus of rats show that sex differences emerging from hormonal signaling can be an important influence on brain structure and function. [11] Work with zebrafish has been especially informative because this species provides the best model for directly linking asymmetric gene expression with asymmetric morphology, and for behavioral analyses. [12]
The left and right hemispheres operate the contralateral sides of the body. Each hemisphere contains sections of all 4 lobes: the frontal lobe, parietal lobe, temporal lobe, and occipital lobe. The two hemispheres are separated along the mediated longitudinal fissure and are connected by the corpus callosum which allows for communication and coordination of stimuli and information. [13] The corpus callosum is the largest collective pathway of white matter tissue in the body that is made of more than 200 million nerve fibers. [14] The left and right hemispheres are associated with different functions and specialize in interpreting the same data in different ways, referred to as lateralization of the brain. The left hemisphere is associated with language and calculations, while the right hemisphere is more closely associated with visual-spatial recognition and facial recognition. This lateralization of brain function results in some specialized regions being only present in a certain hemisphere or being dominant in one hemisphere versus the other. Some of the significant regions included in each hemisphere are listed below. [15]
Some significant regions that can present as asymmetrical in the brain can result in either of the hemispheres due to factors such as genetics. An example would include handedness. Handedness can result from asymmetry in the motor cortex of one hemisphere. For right handed individuals, since the brain operates the contralateral side of the body, they could have a more induced motor cortex in the left hemisphere.
Several diseases have been found to exacerbate brain asymmetries that are already present in the brain. Researchers are starting to look into the effect and relationship of brain asymmetries to diseases such as schizophrenia and dyslexia.
Lateralization of function and asymmetry in the human brain continues to propel a popular branch of neuroscientific and psychological inquiry. Technological advancements for brain mapping have enabled researchers to see more parts of the brain more clearly, which has illuminated previously undetected lateralization differences that occur during different life stages. [9] As more information emerges, researchers are finding insights into how and why early human brains may have evolved the way that they did to adapt to social, environmental and pathological changes. This information provides clues regarding plasticity, or how different parts of the brain can sometimes be recruited for different functions. [26]
Continued study of brain asymmetry also contributes to the understanding and treatment of complex diseases. Neuroimaging in patients with Alzheimer's disease, for example, shows significant deterioration in the left hemisphere, along with a rightward hemispheric dominance—which could relate to recruitment of resources to that side of the brain in the face of damage to the left. [27] These hemispheric changes have been connected to performance on memory tasks. [28]
As has been the case in the past, studies on language processing and the implications of left- and right- handedness also dominate current research on brain asymmetry.
In neuroscience and psychology, the term language center refers collectively to the areas of the brain which serve a particular function for speech processing and production. Language is a core system that gives humans the capacity to solve difficult problems and provides them with a unique type of social interaction. Language allows individuals to attribute symbols to specific concepts, and utilize them through sentences and phrases that follow proper grammatical rules. Finally, speech is the mechanism by which language is orally expressed.
Broca's area, or the Broca area, is a region in the frontal lobe of the dominant hemisphere, usually the left, of the brain with functions linked to speech production.
The vertebrate cerebrum (brain) is formed by two cerebral hemispheres that are separated by a groove, the longitudinal fissure. The brain can thus be described as being divided into left and right cerebral hemispheres. Each of these hemispheres has an outer layer of grey matter, the cerebral cortex, that is supported by an inner layer of white matter. In eutherian (placental) mammals, the hemispheres are linked by the corpus callosum, a very large bundle of nerve fibers. Smaller commissures, including the anterior commissure, the posterior commissure and the fornix, also join the hemispheres and these are also present in other vertebrates. These commissures transfer information between the two hemispheres to coordinate localized functions.
The cingulate cortex is a part of the brain situated in the medial aspect of the cerebral cortex. The cingulate cortex includes the entire cingulate gyrus, which lies immediately above the corpus callosum, and the continuation of this in the cingulate sulcus. The cingulate cortex is usually considered part of the limbic lobe.
The parietal lobe is one of the four major lobes of the cerebral cortex in the brain of mammals. The parietal lobe is positioned above the temporal lobe and behind the frontal lobe and central sulcus.
The temporal lobe is one of the four major lobes of the cerebral cortex in the brain of mammals. The temporal lobe is located beneath the lateral fissure on both cerebral hemispheres of the mammalian brain.
The frontal lobe is the largest of the four major lobes of the brain in mammals, and is located at the front of each cerebral hemisphere. It is parted from the parietal lobe by a groove between tissues called the central sulcus and from the temporal lobe by a deeper groove called the lateral sulcus. The most anterior rounded part of the frontal lobe is known as the frontal pole, one of the three poles of the cerebrum.
The cerebrum, telencephalon or endbrain is the largest part of the brain, containing the cerebral cortex as well as several subcortical structures, including the hippocampus, basal ganglia, and olfactory bulb. In the human brain, the cerebrum is the uppermost region of the central nervous system. The cerebrum develops prenatally from the forebrain (prosencephalon). In mammals, the dorsal telencephalon, or pallium, develops into the cerebral cortex, and the ventral telencephalon, or subpallium, becomes the basal ganglia. The cerebrum is also divided into approximately symmetric left and right cerebral hemispheres.
Brodmann area 45 (BA45), is part of the frontal cortex in the human brain. It is situated on the lateral surface, inferior to BA9 and adjacent to BA46.
Wernicke's area, also called Wernicke's speech area, is one of the two parts of the cerebral cortex that are linked to speech, the other being Broca's area. It is involved in the comprehension of written and spoken language, in contrast to Broca's area, which is primarily involved in the production of language. It is traditionally thought to reside in Brodmann area 22, which is located in the superior temporal gyrus in the dominant cerebral hemisphere, which is the left hemisphere in about 95% of right-handed individuals and 70% of left-handed individuals.
In neuroanatomy, the superior frontal gyrus is a gyrus – a ridge on the brain's cerebral cortex – which makes up about one third of the frontal lobe. It is bounded laterally by the superior frontal sulcus.
In neuroanatomy, the arcuate fasciculus is a bundle of axons that generally connects Broca's area and Wernicke's area in the brain. It is an association fiber tract connecting caudal temporal lobe and inferior frontal lobe.
The planum temporale is the cortical area just posterior to the auditory cortex within the Sylvian fissure. It is a triangular region which forms the heart of Wernicke's area, one of the most important functional areas for language. Original studies on this area found that the planum temporale was one of the most asymmetric regions in the brain, larger in the left cerebral hemisphere than the right.
Brodmann area 22 is a Brodmann's area that is cytoarchitecturally located in the posterior superior temporal gyrus of the brain. In the left cerebral hemisphere, it is one portion of Wernicke's area. The left hemisphere BA22 helps with generation and understanding of individual words. On the right side of the brain, BA22 helps to discriminate pitch and sound intensity, both of which are necessary to perceive melody and prosody. Wernicke's area is active in processing language and consists of the left Brodmann area 22 and Brodmann area 40, the supramarginal gyrus.
The lateralization of brain function is the tendency for some neural functions or cognitive processes to be specialized to one side of the brain or the other. The median longitudinal fissure separates the human brain into two distinct cerebral hemispheres, connected by the corpus callosum. Although the macrostructure of the two hemispheres appears to be almost identical, different composition of neuronal networks allows for specialized function that is different in each hemisphere.
Dichotic listening is a psychological test commonly used to investigate selective attention and the lateralization of brain function within the auditory system. It is used within the fields of cognitive psychology and neuroscience.
Cerebellar cognitive affective syndrome (CCAS), also called Schmahmann's syndrome is a condition that follows from lesions (damage) to the cerebellum of the brain. It refers to a constellation of deficits in the cognitive domains of executive function, spatial cognition, language, and affect resulting from damage to the cerebellum. Impairments of executive function include problems with planning, set-shifting, abstract reasoning, verbal fluency, and working memory, and there is often perseveration, distractibility and inattention. Language problems include dysprosodia, agrammatism and mild anomia. Deficits in spatial cognition produce visual–spatial disorganization and impaired visual–spatial memory. Personality changes manifest as blunting of affect or disinhibited and inappropriate behavior. These cognitive impairments result in an overall lowering of intellectual function. CCAS challenges the traditional view of the cerebellum being responsible solely for regulation of motor functions. It is now thought that the cerebellum is responsible for monitoring both motor and nonmotor functions. The nonmotor deficits described in CCAS are believed to be caused by dysfunction in cerebellar connections to the cerebral cortex and limbic system.
Sign language refers to any natural language which uses visual gestures produced by the hands and body language to express meaning. The brain's left side is the dominant side utilized for producing and understanding sign language, just as it is for speech. In 1861, Paul Broca studied patients with the ability to understand spoken languages but the inability to produce them. The damaged area was named Broca's area, and located in the left hemisphere’s inferior frontal gyrus. Soon after, in 1874, Carl Wernicke studied patients with the reverse deficits: patients could produce spoken language, but could not comprehend it. The damaged area was named Wernicke's area, and is located in the left hemisphere’s posterior superior temporal gyrus.
Disconnection syndrome is a general term for a collection of neurological symptoms caused – via lesions to associational or commissural nerve fibres – by damage to the white matter axons of communication pathways in the cerebrum, independent of any lesions to the cortex. The behavioral effects of such disconnections are relatively predictable in adults. Disconnection syndromes usually reflect circumstances where regions A and B still have their functional specializations except in domains that depend on the interconnections between the two regions.
An estimated 90% of the world's human population consider themselves to be right-handed. The human brain's control of motor function is a mirror image in terms of connectivity; the left hemisphere controls the right hand and vice versa. This theoretically means that the hemisphere contralateral to the dominant hand tends to be more dominant than the ipsilateral hemisphere, however this is not always the case and there are numerous other factors which contribute in complex ways to physical hand preference.