| White matter | |
|---|---|
Micrograph showing white matter with its characteristic fine meshwork-like appearance (left of image - lighter shade of pink) and grey matter, with the characteristic neuronal cell bodies (right of image - dark shade of pink). HPS stain.
| |
Human
brain right dissected lateral view, showing grey matter (the darker
outer parts), and white matter (the inner and prominently whiter parts).
| |
| Details | |
| Location | Central nervous system |
| Identifiers | |
| Latin | substantia alba |
| MeSH | D066127 |
| TA | A14.1.00.009 |
| FMA | 83929 |
White matter structure of human brain (taken by MRI).
White matter refers to areas of the central nervous system (CNS) that are mainly made up of myelinated axons, also called tracts. Long thought to be passive tissue, white matter affects learning and brain functions, modulating the distribution of action potentials, acting as a relay and coordinating communication between different brain regions.
White matter is named for its relatively light appearance resulting from the lipid content of myelin.
However, the tissue of the freshly cut brain appears pinkish-white to
the naked eye because myelin is composed largely of lipid tissue veined
with capillaries. Its white color in prepared specimens is due to its usual preservation in formaldehyde.
Structure
White matter
White matter is composed of bundles, which connect various gray matter areas (the locations of nerve cell bodies) of the brain to each other, and carry nerve impulses between neurons. Myelin acts as an insulator, which allows electrical signals to jump, rather than coursing through the axon, increasing the speed of transmission of all nerve signals.
The total number of long range fibers within a cerebral
hemisphere is 2% of the total number of cortico-cortical fibers (across
cortical areas) and is roughly the same number as those that communicate
between the two hemispheres in the brain's largest white tissue
structure, the corpus callosum. Schüz and Braitenberg note "As a rough rule, the number of fibres of a certain range of lengths is inversely proportional to their length."
White matter in nonelderly adults is 1.7–3.6% blood.
Grey matter
The other main component of the brain is grey matter (actually pinkish tan due to blood capillaries), which is composed of neurons. The substantia nigra is a third colored component found in the brain that appears darker due to higher levels of melanin in dopaminergic neurons than its nearby areas. Note that white matter can sometimes appear darker than grey matter on a microscope slide because of the type of stain used. Cerebral- and spinal white matter do not contain dendrites, neural cell bodies, or shorter axons, which can only be found in grey matter.
Location
White matter forms the bulk of the deep parts of the brain and the superficial parts of the spinal cord. Aggregates of grey matter such as the basal ganglia (caudate nucleus, putamen, globus pallidus, substantia nigra, subthalamic nucleus, nucleus accumbens) and brainstem nuclei (red nucleus, cranial nerve nuclei) are spread within the cerebral white matter.
The cerebellum
is structured in a similar manner as the cerebrum, with a superficial
mantle of cerebellar cortex, deep cerebellar white matter (called the "arbor vitae") and aggregates of grey matter surrounded by deep cerebellar white matter (dentate nucleus, globose nucleus, emboliform nucleus, and fastigial nucleus). The fluid-filled cerebral ventricles (lateral ventricles, third ventricle, cerebral aqueduct, fourth ventricle) are also located deep within the cerebral white matter.
Myelinated axon length
Men
have more white matter than women both in volume and in length of
myelinated axons. At the age of 20, the total length of myelinated
fibers in men is 176,000 km while that of a woman is 149,000 km.
There is a decline in total length with age of about 10% each decade
such that a man at 80 years of age has 97,200 km and a female 82,000 km. Most of this reduction is due to the loss of thinner fibers.
Function
White
matter is the tissue through which messages pass between different
areas of gray matter within the central nervous system. The white matter
is white because of the fatty substance (myelin) that surrounds the
nerve fibers (axons). This myelin is found in almost all long nerve
fibers, and acts as an electrical insulation. This is important because
it allows the messages to pass quickly from place to place.
Unlike gray matter, which peaks in development in a person's
twenties, the white matter continues to develop, and peaks in middle
age.
Research
Multiple sclerosis (MS) is the most common of the inflammatory demyelinating diseases of the central nervous system which affect white matter. In MS lesions, the myelin sheath around the axons is deteriorated by inflammation. Alcohol use disorders are associated with a decrease in white matter volume.
Amyloid plaques in white matter may be associated with Alzheimer's disease and other neurodegenerative diseases. Other changes that commonly occur with age include the development of leukoaraiosis,
which is a rarefaction of the white matter that can be correlated with a
variety of conditions, including loss of myelin pallor, axonal loss,
and diminished restrictive function of the blood–brain barrier.
White matter lesions on magnetic resonance imaging are linked to several adverse outcomes, such as cognitive impairment and depression. White matter hyperintensity are more than often present with vascular dementia, particularly among small vessel/subcortical subtypes of vascular dementia.
Volume
Smaller volumes (in terms of group averages) of white matter might be associated with larger deficits in attention, declarative memory, executive functions, intelligence, and academic achievement. However, volume change is continuous throughout one's lifetime due to neuroplasticity,
and is a contributing factor rather than determinant factor of certain
functional deficits due to compensating effects in other brain regions. The integrity of white matter declines due to aging. Nonetheless, regular aerobic exercise appears to either postpone the aging effect or in turn enhance the white matter integrity in the long run. Changes in white matter volume due to inflammation or injury may be a factor in the severity of obstructive sleep apnea.
Imaging
The study of white matter has been advanced with the neuroimaging technique called diffusion tensor imaging where magnetic resonance imaging (MRI) brain scanners are used. As of 2007, more than 700 publications have been published on the subject.
A 2009 paper by Jan Scholz and colleagues
used diffusion tensor imaging (DTI) to demonstrate changes in white
matter volume as a result of learning a new motor task (e.g. juggling).
The study is important as the first paper to correlate motor learning
with white matter changes. Previously, many researchers had considered
this type of learning to be exclusively mediated by dendrites, which are
not present in white matter. The authors suggest that electrical
activity in axons may regulate myelination in axons. Or, gross changes
in the diameter or packing density of the axon might cause the change.
A more recent DTI study by Sampaio-Baptista and colleagues reported
changes in white matter with motor learning along with increases in
myelination.
