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From Wikipedia, the free encyclopedia
This article is about the brains of all types of animals, including humans. For
information specific to the human brain, see Human brain. For other uses, see Brain
(disambiguation) and Brains (disambiguation).
Not to be confused with Brian, Braine, or Brane.
Brain
The brain of a chimpanzee
Identifiers
MeSH D001921
NeuroNames 21
TA98 A14.1.03.001
TA2 5415
Anatomical terminology
[edit on Wikidata]

The brain (or encephalon) is an organ that serves as the center of the nervous
system in all vertebrate and most invertebrate animals. The brain is the largest
cluster of neurons in the body and is typically located in the head, usually near
organs for special senses such as vision, hearing and olfaction. It is the most
specialized and energy-consuming organ in the body, responsible for complex sensory
perception, motor control, endocrine regulation and the development of
intelligence.

While invertebrate brains arise from paired segmental ganglia (each of which is
only responsible for the respective body segment) of the ventral nerve cord,
vertebrate brains develop axially from the midline dorsal nerve cord as a vesicular
enlargement at the rostral end of the neural tube, with centralized control over
all body segments. All vertebrate brains can be embryonically divided into three
parts: the forebrain (prosencephalon, subdivided into telencephalon and
diencephalon), midbrain (mesencephalon) and hindbrain (rhombencephalon, subdivided
into metencephalon and myelencephalon). The spinal cord, which directly interacts
with somatic functions below the head, can be considered a caudal extension of the
myelencephalon enclosed inside the vertebral column. Together, the brain and spinal
cord constitute the central nervous system in all vertebrates.

In humans, the cerebral cortex contains approximately 14–16 billion neurons,[1] and
the estimated number of neurons in the cerebellum is 55–70 billion.[2] Each neuron
is connected by synapses to several thousand other neurons, typically communicating
with one another via root-like protrusions called dendrites and long fiber-like
extensions called axons, which are usually myelinated and carry trains of rapid
micro-electric signal pulses called action potentials to target specific recipient
cells in other areas of the brain or distant parts of the body. The prefrontal
cortex, which controls executive functions, is particularly well developed in
humans.

Physiologically, brains exert centralized control over a body's other organs. They
act on the rest of the body both by generating patterns of muscle activity and by
driving the secretion of chemicals called hormones. This centralized control allows
rapid and coordinated responses to changes in the environment. Some basic types of
responsiveness such as reflexes can be mediated by the spinal cord or peripheral
ganglia, but sophisticated purposeful control of behavior based on complex sensory
input requires the information integrating capabilities of a centralized brain.

The operations of individual brain cells are now understood in considerable detail
but the way they cooperate in ensembles of millions is yet to be solved.[3] Recent
models in modern neuroscience treat the brain as a biological computer, very
different in mechanism from a digital computer, but similar in the sense that it
acquires information from the surrounding world, stores it, and processes it in a
variety of ways.

This article compares the properties of brains across the entire range of animal
species, with the greatest attention to vertebrates. It deals with the human brain
insofar as it shares the properties of other brains. The ways in which the human
brain differs from other brains are covered in the human brain article. Several
topics that might be covered here are instead covered there because much more can
be said about them in a human context. The most important that are covered in the
human brain article are brain disease and the effects of brain damage.
Anatomy
a blob with a blue patch in the center, surrounded by a white area, surrounded by a
thin strip of dark-colored material
Cross section of the olfactory bulb of a rat, stained in two different ways at the
same time: one stain shows neuron cell bodies, the other shows receptors for the
neurotransmitter GABA.

The shape and size of the brain varies greatly between species, and identifying
common features is often difficult.[4] Nevertheless, there are a number of
principles of brain architecture that apply across a wide range of species.[5] Some
aspects of brain structure are common to almost the entire range of animal species;
[6] others distinguish "advanced" brains from more primitive ones, or distinguish
vertebrates from invertebrates.[4]

The simplest way to gain information about brain anatomy is by visual inspection,
but many more sophisticated techniques have been developed. Brain tissue in its
natural state is too soft to work with, but it can be hardened by immersion in
alcohol or other fixatives, and then sliced apart for examination of the interior.
Visually, the interior of the brain consists of areas of so-called grey matter,
with a dark color, separated by areas of white matter, with a lighter color.
Further information can be gained by staining slices of brain tissue with a variety
of chemicals that bring out areas where specific types of molecules are present in
high concentrations. It is also possible to examine the microstructure of brain
tissue using a microscope, and to trace the pattern of connections from one brain
area to another.[7]
Cellular structure
drawing showing a neuron with a fiber emanating from it labeled "axon" and making
contact with another cell. An inset shows an enlargement of the contact zone.
Neurons generate electrical signals that travel along their axons. When a pulse of
electricity reaches a junction called a synapse, it causes a neurotransmitter
chemical to be released, which binds to receptors on other cells and thereby alters
their electrical activity.

The brains of all species are composed primarily of two broad classes of cells:
neurons and glial cells. Glial cells (also known as glia or neuroglia) come in
several types, and perform a number of critical functions, including structural
support, metabolic support, insulation, and guidance of development. Neurons,
however, are usually considered the most important cells in the brain.[8] The
property that makes neurons unique is their ability to send signals to specific
target cells over long distances.[8] They send these signals by means of an axon,
which is a thin protoplasmic fiber that extends from the cell body and projects,
usually with numerous branches, to other areas, sometimes nearby, sometimes in
distant parts of the brain or body. The length of an axon can be extraordinary: for
example, if a pyramidal cell (an excitatory neuron) of the cerebral cortex were
magnified so that its cell body became the size of a human body, its axon, equally
magnified, would become a cable a few centimeters in diameter, extending more than
a kilometer.[9] These axons transmit signals in the form of electrochemical pulses
called action potentials, which last less than a thousandth of a second and travel
along the axon at speeds of 1–100 meters per second. Some neurons emit action
potentials constantly, at rates of 10–100 per second, usually in irregular
patterns; other neurons are quiet most of the time, but occasionally emit a burst
of action potentials.[10]

Axons transmit signals to other neurons by means of specialized junctions called

synapses. A single axon may make as many as several thousand synaptic connections
with other cells.[8] When an action potential, traveling along an axon, arrives at
a synapse, it causes a chemical called a neurotransmitter to be released. The
neurotransmitter binds to receptor molecules in the membrane of the target cell.[8]

Synapses are the key functional elements of the brain.[11] The essential function
of the brain is cell-to-cell communication, and synapses are the points at which
communication occurs. The human brain has been estimated to contain approximately
100 trillion synapses;[12] even the brain of a fruit fly contains several million.
[13] The functions of these synapses are very diverse: some are excitatory
(exciting the target cell); others are inhibitory; others work by activating second
messenger systems that change the internal chemistry of their target cells in
complex ways.[11] A large number of synapses are dynamically modifiable; that is,
they are capable of changing strength in a way that is controlled by the patterns
of signals that pass through them. It is widely believed that activity-dependent
modification of synapses is the brain's primary mechanism for learning and memory.
[11]

Most of the space in the brain is taken up by axons, which are often bundled
together in what are called nerve fiber tracts. A myelinated axon is wrapped in a
fatty insulating sheath of myelin, which serves to greatly increase the speed of
signal propagation. (There are also unmyelinated axons). Myelin is white, making
parts of the brain filled exclusively with nerve fibers appear as light-colored
white matter, in contrast to the darker-colored grey matter that marks areas with
high densities of neuron cell bodies.[8]
Evolution
Main article: Evolution of the brain
Generic bilaterian nervous system
A rod-shaped body contains a digestive system running from the mouth at one end to
the anus at the other. Alongside the digestive system is a nerve cord with a brain
at the end, near to the mouth.
Nervous system of a generic bilaterian animal, in the form of a nerve cord with
segmental enlargements, and a "brain" at the front

Except for a few primitive organisms such as sponges (which have no nervous system)
[14] and cnidarians (which have a diffuse nervous system consisting of a nerve
net),[14] all living multicellular animals are bilaterians, meaning animals with a
bilaterally symmetric body plan (that is, left and right sides that are approximate
mirror images of each other).[15] All bilaterians are thought to have descended
from a common ancestor that appeared late in the Cryogenian period, 700–650 million
years ago, and it has been hypothesized that this common ancestor had the shape of
a simple tubeworm with a segmented body.[15] At a schematic level, that basic worm-
shape continues to be reflected in the body and nervous system architecture of all
modern bilaterians, including vertebrates.[16] The fundamental bilateral body form
is a tube with a hollow gut cavity running from the mouth to the anus, and a nerve
cord with an enlargement (a ganglion) for each body segment, with an especially
large ganglion at the front, called the brain. The brain is small and simple in
some species, such as nematode worms; in other species, including vertebrates, it
is the most complex organ in the body.[4] Some types of worms, such as leeches,
also have an enlarged ganglion at the back end of the nerve cord, known as a "tail
brain".[17]

There are a few types of existing bilaterians that lack a recognizable brain,
including echinoderms and tunicates. It has not been definitively established
whether the existence of these brainless species indicates that the earliest
bilaterians lacked a brain, or whether their ancestors evolved in a way that led to
the disappearance of a previously existing brain structure.
Invertebrates
A fly resting on a reflective surface. A large, red eye faces the camera. The body
appears transparent, apart from black pigment at the end of its abdomen.
Fruit flies (Drosophila) have been extensively studied to gain insight into the
role of genes in brain development.

This category includes tardigrades, arthropods, molluscs, and numerous types of

worms. The diversity of invertebrate body plans is matched by an equal diversity in
brain structures.[18]

Two groups of invertebrates have notably complex brains: arthropods (insects,

crustaceans, arachnids, and others), and cephalopods (octopuses, squids, and
similar molluscs).[19] The brains of arthropods and cephalopods arise from twin
parallel nerve cords that extend through the body of the animal. Arthropods have a
central brain, the supraesophageal ganglion, with three divisions and large optical
lobes behind each eye for visual processing.[19] Cephalopods such as the octopus
and squid have the largest brains of any invertebrates.[20]
There are several invertebrate species whose brains have been studied intensively
because they have properties that make them convenient for experimental work:

Fruit flies (Drosophila), because of the large array of techniques available

for studying their genetics, have been a natural subject for studying the role of
genes in brain development.[21] In spite of the large evolutionary distance between
insects and mammals, many aspects of Drosophila neurogenetics have been shown to be
relevant to humans. The first biological clock genes, for example, were identified
by examining Drosophila mutants that showed disrupted daily activity cycles.[22] A
search in the genomes of vertebrates revealed a set of analogous genes, which were
found to play similar roles in the mouse biological clock—and therefore almost
certainly in the human biological clock as well.[23] Studies done on Drosophila,
also show that most neuropil regions of the brain are continuously reorganized
throughout life in response to specific living conditions.[24]
The nematode worm Caenorhabditis elegans, like Drosophila, has been studied
largely because of its importance in genetics.[25] In the early 1970s, Sydney
Brenner chose it as a model organism for studying the way that genes control
development. One of the advantages of working with this worm is that the body plan
is very stereotyped: the nervous system of the hermaphrodite contains exactly 302
neurons, always in the same places, making identical synaptic connections in every
worm.[26] Brenner's team sliced worms into thousands of ultrathin sections and
photographed each one under an electron microscope, then visually matched fibers
from section to section, to map out every neuron and synapse in the entire body.
[27] The complete neuronal wiring diagram of C.elegans – its connectome was
achieved.[28] Nothing approaching this level of detail is available for any other
organism, and the information gained has enabled a multitude of studies that would
otherwise have not been possible.[29]
The sea slug Aplysia californica was chosen by Nobel Prize-winning
neurophysiologist Eric Kandel as a model for studying the cellular basis of
learning and memory, because of the simplicity and accessibility of its nervous
system, and it has been examined in hundreds of experiments.[30]

Vertebrates
A T-shaped object is made up of the cord at the bottom which feeds into a lower
central mass. This is topped by a larger central mass with an arm extending from
either side.
The brain of a shark

The first vertebrates appeared over 500 million years ago (Mya), during the
Cambrian period, and may have resembled the modern hagfish in form.[31] Jawed fish
appeared by 445 Mya, amphibians by 350 Mya, reptiles by 310 Mya and mammals by 200
Mya (approximately). Each species has an equally long evolutionary history, but the
brains of modern hagfishes, lampreys, sharks, amphibians, reptiles, and mammals
show a gradient of size and complexity that roughly follows the evolutionary
sequence. All of these brains contain the same set of basic anatomical components,
but many are rudimentary in the hagfish, whereas in mammals the foremost part (the
telencephalon) is greatly elaborated and expanded.[32]

Brains are most commonly compared in terms of their size. The relationship between
brain size, body size and other variables has been studied across a wide range of
vertebrate species. As a rule, brain size increases with body size, but not in a
simple linear proportion. In general, smaller animals tend to have larger brains,
measured as a fraction of body size. For mammals, the relationship between brain
volume and body mass essentially follows a power law with an exponent of about
0.75.[33] This formula describes the central tendency, but every family of mammals
departs from it to some degree, in a way that reflects in part the complexity of
their behavior. For example, primates have brains 5 to 10 times larger than the
formula predicts. Predators tend to have larger brains than their prey, relative to
body size.[34]
The nervous system is shown as a rod with protrusions along its length. The spinal
cord at the bottom connects to the hindbrain which widens out before narrowing
again. This is connected to the midbrain, which again bulges, and which finally
connects to the forebrain which has two large protrusions.
The main subdivisions of the embryonic vertebrate brain (left), which later
differentiate into structures of the adult brain (right)

All vertebrate brains share a common underlying form, which appears most clearly
during early stages of embryonic development. In its earliest form, the brain
appears as three swellings at the front end of the neural tube; these swellings
eventually become the forebrain, midbrain, and hindbrain (the prosencephalon,
mesencephalon, and rhombencephalon, respectively). At the earliest stages of brain
development, the three areas are roughly equal in size. In many classes of
vertebrates, such as fish and amphibians, the three parts remain similar in size in
the adult, but in mammals the forebrain becomes much larger than the other parts,
and the midbrain becomes very small.[8]

The brains of vertebrates are made of very soft tissue.[8] Living brain tissue is
pinkish on the outside and mostly white on the inside, with subtle variations in
color. Vertebrate brains are surrounded by a system of connective tissue membranes
called meninges that separate the skull from the brain. Blood vessels enter the
central nervous system through holes in the meningeal layers. The cells in the
blood vessel walls are joined tightly to one another, forming the blood–brain
barrier, which blocks the passage of many toxins and pathogens[35] (though at the
same time blocking antibodies and some drugs, thereby presenting special challenges
in treatment of diseases of the brain).[36]

Neuroanatomists usually divide the vertebrate brain into six main regions: the
telencephalon (cerebral hemispheres), diencephalon (thalamus and hypothalamus),
mesencephalon (midbrain), cerebellum, pons, and medulla oblongata. Each of these
areas has a complex internal structure. Some parts, such as the cerebral cortex and
the cerebellar cortex, consist of layers that are folded or convoluted to fit
within the available space. Other parts, such as the thalamus and hypothalamus,
consist of clusters of many small nuclei. Thousands of distinguishable areas can be
identified within the vertebrate brain based on fine distinctions of neural
structure, chemistry, and connectivity.[8]
Corresponding regions of human and shark brain are shown. The shark brain is
splayed out, while the human brain is more compact. The shark brain starts with the
medulla, which is surrounded by various structures, and ends with the
telencephalon. The cross-section of the human brain shows the medulla at the bottom
surrounded by the same structures, with the telencephalon thickly coating the top
of the brain.
The main anatomical regions of the vertebrate brain, shown for shark and human. The
same parts are present, but they differ greatly in size and shape.

Although the same basic components are present in all vertebrate brains, some
branches of vertebrate evolution have led to substantial distortions of brain
geometry, especially in the forebrain area. The brain of a shark shows the basic
components in a straightforward way, but in teleost fishes (the great majority of
existing fish species), the forebrain has become "everted", like a sock turned
inside out. In birds, there are also major changes in forebrain structure.[37]
These distortions can make it difficult to match brain components from one species
with those of another species.[38]

Here is a list of some of the most important vertebrate brain components, along
with a brief description of their functions as currently understood:
See also: List of regions in the human brain
 The medulla, along with the spinal cord, contains many small nuclei involved in
a wide variety of sensory and involuntary motor functions such as vomiting, heart
rate and digestive processes.[8]
The pons lies in the brainstem directly above the medulla. Among other things,
it contains nuclei that control often voluntary but simple acts such as sleep,
respiration, swallowing, bladder function, equilibrium, eye movement, facial
expressions, and posture.[39]
The hypothalamus is a small region at the base of the forebrain, whose
complexity and importance belies its size. It is composed of numerous small nuclei,
each with distinct connections and neurochemistry. The hypothalamus is engaged in
additional involuntary or partially voluntary acts such as sleep and wake cycles,
eating and drinking, and the release of some hormones.[40]
The thalamus is a collection of nuclei with diverse functions: some are
involved in relaying information to and from the cerebral hemispheres, while others
are involved in motivation. The subthalamic area (zona incerta) seems to contain
action-generating systems for several types of "consummatory" behaviors such as
eating, drinking, defecation, and copulation.[41]
The cerebellum modulates the outputs of other brain systems, whether motor-
related or thought related, to make them certain and precise. Removal of the
cerebellum does not prevent an animal from doing anything in particular, but it
makes actions hesitant and clumsy. This precision is not built-in but learned by
trial and error. The muscle coordination learned while riding a bicycle is an
example of a type of neural plasticity that may take place largely within the
cerebellum.[8] 10% of the brain's total volume consists of the cerebellum and 50%
of all neurons are held within its structure.[42]
The optic tectum allows actions to be directed toward points in space, most
commonly in response to visual input. In mammals, it is usually referred to as the
superior colliculus, and its best-studied function is to direct eye movements. It
also directs reaching movements and other object-directed actions. It receives
strong visual inputs, but also inputs from other senses that are useful in
directing actions, such as auditory input in owls and input from the
thermosensitive pit organs in snakes. In some primitive fishes, such as lampreys,
this region is the largest part of the brain.[43] The superior colliculus is part
of the midbrain.
The pallium is a layer of grey matter that lies on the surface of the forebrain
and is the most complex and most recent evolutionary development of the brain as an
organ.[44] In reptiles and mammals, it is called the cerebral cortex. Multiple
functions involve the pallium, including smell and spatial memory. In mammals,
where it becomes so large as to dominate the brain, it takes over functions from
many other brain areas. In many mammals, the cerebral cortex consists of folded
bulges called gyri that create deep furrows or fissures called sulci. The folds
increase the surface area of the cortex and therefore increase the amount of gray
matter and the amount of information that can be stored and processed.[45]
The hippocampus, strictly speaking, is found only in mammals. However, the area
it derives from, the medial pallium, has counterparts in all vertebrates. There is
evidence that this part of the brain is involved in complex events such as spatial
memory and navigation in fishes, birds, reptiles, and mammals.[46]
The basal ganglia are a group of interconnected structures in the forebrain.
The primary function of the basal ganglia appears to be action selection: they send
inhibitory signals to all parts of the brain that can generate motor behaviors, and
in the right circumstances can release the inhibition, so that the action-
generating systems are able to execute their actions. Reward and punishment exert
their most important neural effects by altering connections within the basal
ganglia.[47]
The olfactory bulb is a special structure that processes olfactory sensory
signals and sends its output to the olfactory part of the pallium. It is a major
brain component in many vertebrates, but is greatly reduced in humans and other
primates (whose senses are dominated by information acquired by sight rather than
smell).[48]
Birds
Main article: Avian brain
This section is an excerpt from Avian brain.[edit]
Brains of an emu, a kiwi, a barn owl, and a pigeon, with visual processing areas
labelled

The avian brain is the central organ of the nervous system in birds. Birds possess
large, complex brains, which process, integrate, and coordinate information
received from the environment and make decisions on how to respond with the rest of
the body. Like in all chordates, the avian brain is contained within the skull
bones of the head.

The bird brain is divided into a number of sections, each with a different
function. The cerebrum or telencephalon is divided into two hemispheres, and
controls higher functions. The telencephalon is dominated by a large pallium, which
corresponds to the mammalian cerebral cortex and is responsible for the cognitive
functions of birds. The pallium is made up of several major structures: the
hyperpallium, a dorsal bulge of the pallium found only in birds, as well as the
nidopallium, mesopallium, and archipallium. The bird telencephalon nuclear
structure, wherein neurons are distributed in three-dimensionally arranged
clusters, with no large-scale separation of white matter and grey matter, though
there exist layer-like and column-like connections. Structures in the pallium are
associated with perception, learning, and cognition. Beneath the pallium are the
two components of the subpallium, the striatum and pallidum. The subpallium
connects different parts of the telencephalon and plays major roles in a number of
critical behaviours. To the rear of the telencephalon are the thalamus, midbrain,
and cerebellum. The hindbrain connects the rest of the brain to the spinal cord.
The size and structure of the avian brain enables prominent behaviours of birds
such as flight and vocalization. Dedicated structures and pathways integrate the
auditory and visual senses, strong in most species of birds, as well as the
typically weaker olfactory and tactile senses. Social behaviour, widespread among
birds, depends on the organisation and functions of the brain. Some birds exhibit
strong abilities of cognition, enabled by the unique structure and physiology of
the avian brain.
Mammals

The most obvious difference between the brains of mammals and other vertebrates is
in terms of size. On average, a mammal has a brain roughly twice as large as that
of a bird of the same body size, and ten times as large as that of a reptile of the
same body size.[49]

Size, however, is not the only difference: there are also substantial differences
in shape. The hindbrain and midbrain of mammals are generally similar to those of
other vertebrates, but dramatic differences appear in the forebrain, which is
greatly enlarged and also altered in structure.[50] The cerebral cortex is the part
of the brain that most strongly distinguishes mammals. In non-mammalian
vertebrates, the surface of the cerebrum is lined with a comparatively simple
three-layered structure called the pallium. In mammals, the pallium evolves into a
complex six-layered structure called neocortex or isocortex.[51] Several areas at
the edge of the neocortex, including the hippocampus and amygdala, are also much
more extensively developed in mammals than in other vertebrates.[50]

The elaboration of the cerebral cortex carries with it changes to other brain
areas. The superior colliculus, which plays a major role in visual control of
behavior in most vertebrates, shrinks to a small size in mammals, and many of its
functions are taken over by visual areas of the cerebral cortex.[49] The cerebellum
of mammals contains a large portion (the neocerebellum) dedicated to supporting the
cerebral cortex, which has no counterpart in other vertebrates.[52]
Primates
See also: Human brain
Encephalization Quotient Species EQ[53]
Human 7.4–7.8
Common chimpanzee 2.2–2.5
Rhesus monkey 2.1
Bottlenose dolphin 4.14[54]
Elephant 1.13–2.36[55]
Dog 1.2
Horse 0.9
Rat 0.4

The brains of humans and other primates contain the same structures as the brains
of other mammals, but are generally larger in proportion to body size.[56] The
encephalization quotient (EQ) is used to compare brain sizes across species. It
takes into account the nonlinearity of the brain-to-body relationship.[53] Humans
have an average EQ in the 7-to-8 range, while most other primates have an EQ in the
2-to-3 range. Dolphins have values higher than those of primates other than humans,
[54] but nearly all other mammals have EQ values that are substantially lower.

Most of the enlargement of the primate brain comes from a massive expansion of the
cerebral cortex, especially the prefrontal cortex and the parts of the cortex
involved in vision.[57] The visual processing network of primates includes at least
30 distinguishable brain areas, with a complex web of interconnections. It has been
estimated that visual processing areas occupy more than half of the total surface
of the primate neocortex.[58] The prefrontal cortex carries out functions that
include planning, working memory, motivation, attention, and executive control. It
takes up a much larger proportion of the brain for primates than for other species,
and an especially large fraction of the human brain.[59]
Development
Main article: Neural development
Very simple drawing of the front end of a human embryo, showing each vesicle of the
developing brain in a different color.
Brain of a human embryo in the sixth week of development

The brain develops in an intricately orchestrated sequence of stages.[60] It

changes in shape from a simple swelling at the front of the nerve cord in the
earliest embryonic stages, to a complex array of areas and connections. Neurons are
created in special zones that contain stem cells, and then migrate through the
tissue to reach their ultimate locations. Once neurons have positioned themselves,
their axons sprout and navigate through the brain, branching and extending as they
go, until the tips reach their targets and form synaptic connections. In a number
of parts of the nervous system, neurons and synapses are produced in excessive
numbers during the early stages, and then the unneeded ones are pruned away.[60]

For vertebrates, the early stages of neural development are similar across all
species.[60] As the embryo transforms from a round blob of cells into a wormlike
structure, a narrow strip of ectoderm running along the midline of the back is
induced to become the neural plate, the precursor of the nervous system. The neural
plate folds inward to form the neural groove, and then the lips that line the
groove merge to enclose the neural tube, a hollow cord of cells with a fluid-filled
ventricle at the center. At the front end, the ventricles and cord swell to form
three vesicles that are the precursors of the prosencephalon (forebrain),
mesencephalon (midbrain), and rhombencephalon (hindbrain). At the next stage, the
forebrain splits into two vesicles called the telencephalon (which will contain the
cerebral cortex, basal ganglia, and related structures) and the diencephalon (which
will contain the thalamus and hypothalamus). At about the same time, the hindbrain
splits into the metencephalon (which will contain the cerebellum and pons) and the
myelencephalon (which will contain the medulla oblongata). Each of these areas
contains proliferative zones where neurons and glial cells are generated; the
resulting cells then migrate, sometimes for long distances, to their final
positions.[60]

Once a neuron is in place, it extends dendrites and an axon into the area around
it. Axons, because they commonly extend a great distance from the cell body and
need to reach specific targets, grow in a particularly complex way. The tip of a
growing axon consists of a blob of protoplasm called a growth cone, studded with
chemical receptors. These receptors sense the local environment, causing the growth
cone to be attracted or repelled by various cellular elements, and thus to be
pulled in a particular direction at each point along its path. The result of this
pathfinding process is that the growth cone navigates through the brain until it
reaches its destination area, where other chemical cues cause it to begin
generating synapses. Considering the entire brain, thousands of genes create
products that influence axonal pathfinding.[60]

The synaptic network that finally emerges is only partly determined by genes,
though. In many parts of the brain, axons initially "overgrow", and then are
"pruned" by mechanisms that depend on neural activity.[60] In the projection from
the eye to the midbrain, for example, the structure in the adult contains a very
precise mapping, connecting each point on the surface of the retina to a
corresponding point in a midbrain layer. In the first stages of development, each
axon from the retina is guided to the right general vicinity in the midbrain by
chemical cues, but then branches very profusely and makes initial contact with a
wide swath of midbrain neurons. The retina, before birth, contains special
mechanisms that cause it to generate waves of activity that originate spontaneously
at a random point and then propagate slowly across the retinal layer. These waves
are useful because they cause neighboring neurons to be active at the same time;
that is, they produce a neural activity pattern that contains information about the
spatial arrangement of the neurons. This information is exploited in the midbrain
by a mechanism that causes synapses to weaken, and eventually vanish, if activity
in an axon is not followed by activity of the target cell. The result of this
sophisticated process is a gradual tuning and tightening of the map, leaving it
finally in its precise adult form.[61]

Similar things happen in other brain areas: an initial synaptic matrix is generated
as a result of genetically determined chemical guidance, but then gradually refined
by activity-dependent mechanisms, partly driven by internal dynamics, partly by
external sensory inputs. In some cases, as with the retina-midbrain system,
activity patterns depend on mechanisms that operate only in the developing brain,
and apparently exist solely to guide development.[61]

In humans and many other mammals, new neurons are created mainly before birth, and
the infant brain contains substantially more neurons than the adult brain.[60]
There are, however, a few areas where new neurons continue to be generated
throughout life. The two areas for which adult neurogenesis is well established are
the olfactory bulb, which is involved in the sense of smell, and the dentate gyrus
of the hippocampus, where there is evidence that the new neurons play a role in
storing newly acquired memories. With these exceptions, however, the set of neurons
that is present in early childhood is the set that is present for life. Glial cells
are different: as with most types of cells in the body, they are generated
throughout the lifespan.[62]

There has long been debate about whether the qualities of mind, personality, and
intelligence can be attributed to heredity or to upbringing—this is the nature and
nurture controversy.[63] Although many details remain to be settled, neuroscience
research has clearly shown that both factors are important. Genes determine the
general form of the brain, and genes determine how the brain reacts to experience.
Experience, however, is required to refine the matrix of synaptic connections,
which in its developed form contains far more information than the genome does. In
some respects, all that matters is the presence or absence of experience during
critical periods of development.[64] In other respects, the quantity and quality of
experience are important; for example, there is substantial evidence that animals
raised in enriched environments have thicker cerebral cortices, indicating a higher
density of synaptic connections, than animals whose levels of stimulation are
restricted.[65]
Physiology

The functions of the brain depend on the ability of neurons to transmit

electrochemical signals to other cells, and their ability to respond appropriately
to electrochemical signals received from other cells. The electrical properties of
neurons are controlled by a wide variety of biochemical and metabolic processes,
most notably the interactions between neurotransmitters and receptors that take
place at synapses.[8]
Neurotransmitters and receptors

Neurotransmitters are chemicals that are released at synapses when the local
membrane is depolarised and Ca2+ enters into the cell, typically when an action
potential arrives at the synapse – neurotransmitters attach themselves to receptor
molecules on the membrane of the synapse's target cell (or cells), and thereby
alter the electrical or chemical properties of the receptor molecules. With few
exceptions, each neuron in the brain releases the same chemical neurotransmitter,
or combination of neurotransmitters, at all the synaptic connections it makes with
other neurons; this rule is known as Dale's principle.[8] Thus, a neuron can be
characterized by the neurotransmitters that it releases. The great majority of
psychoactive drugs exert their effects by altering specific neurotransmitter
systems. This applies to drugs such as cannabinoids, nicotine, heroin, cocaine,
alcohol, fluoxetine, chlorpromazine, and many others.[66]

The two neurotransmitters that are most widely found in the vertebrate brain are
glutamate, which almost always exerts excitatory effects on target neurons, and
gamma-aminobutyric acid (GABA), which is almost always inhibitory. Neurons using
these transmitters can be found in nearly every part of the brain.[67] Because of
their ubiquity, drugs that act on glutamate or GABA tend to have broad and powerful
effects. Some general anesthetics act by reducing the effects of glutamate; most
tranquilizers exert their sedative effects by enhancing the effects of GABA.[68]

There are dozens of other chemical neurotransmitters that are used in more limited
areas of the brain, often areas dedicated to a particular function. Serotonin, for
example—the primary target of many antidepressant drugs and many dietary aids—comes
exclusively from a small brainstem area called the raphe nuclei.[69]
Norepinephrine, which is involved in arousal, comes exclusively from a nearby small
area called the locus coeruleus.[70] Other neurotransmitters such as acetylcholine
and dopamine have multiple sources in the brain but are not as ubiquitously
distributed as glutamate and GABA.[71]
Electrical activity
Graph showing 16 voltage traces going across the page from left to right, each
showing a different signal. At the middle of the page all of the traces abruptly
begin to show sharp jerky spikes, which continue to the end of the plot.
Brain electrical activity recorded from a human patient during an epileptic seizure

As a side effect of the electrochemical processes used by neurons for signaling,

brain tissue generates electric fields when it is active. When large numbers of
neurons show synchronized activity, the electric fields that they generate can be
large enough to detect outside the skull, using electroencephalography (EEG)[72] or
magnetoencephalography (MEG). EEG recordings, along with recordings made from
electrodes implanted inside the brains of animals such as rats, show that the brain
of a living animal is constantly active, even during sleep.[73] Each part of the
brain shows a mixture of rhythmic and nonrhythmic activity, which may vary
according to behavioral state. In mammals, the cerebral cortex tends to show large
slow delta waves during sleep, faster alpha waves when the animal is awake but
inattentive, and chaotic-looking irregular activity when the animal is actively
engaged in a task, called beta and gamma waves. During an epileptic seizure, the
brain's inhibitory control mechanisms fail to function and electrical activity
rises to pathological levels, producing EEG traces that show large wave and spike
patterns not seen in a healthy brain. Relating these population-level patterns to
the computational functions of individual neurons is a major focus of current
research in neurophysiology.[73]
Metabolism

All vertebrates have a blood–brain barrier that allows metabolism inside the brain
to operate differently from metabolism in other parts of the body. The
neurovascular unit regulates cerebral blood flow so that activated neurons can be
supplied with energy. Glial cells play a major role in brain metabolism by
controlling the chemical composition of the fluid that surrounds neurons, including
levels of ions and nutrients.[74]

Brain tissue consumes a large amount of energy in proportion to its volume, so

large brains place severe metabolic demands on animals. The need to limit body
weight in order, for example, to fly, has apparently led to selection for a
reduction of brain size in some species, such as bats.[75] Most of the brain's
energy consumption goes into sustaining the electric charge (membrane potential) of
neurons.[74] Most vertebrate species devote between 2% and 8% of basal metabolism
to the brain. In primates, however, the percentage is much higher—in humans it
rises to 20–25%.[76] The energy consumption of the brain does not vary greatly over
time, but active regions of the cerebral cortex consume somewhat more energy than
inactive regions; this forms the basis for the functional brain imaging methods of
PET, fMRI,[77] and NIRS.[78] The brain typically gets most of its energy from
oxygen-dependent metabolism of glucose (i.e., blood sugar),[74] but ketones provide
a major alternative source, together with contributions from medium chain fatty
acids (caprylic and heptanoic acids),[79][80] lactate,[81] acetate,[82] and
possibly amino acids.[83]
Function
Model of a neural circuit in the cerebellum, as proposed by James S. Albus

Information from the sense organs is collected in the brain. There it is used to
determine what actions the organism is to take. The brain processes the raw data to
extract information about the structure of the environment. Next it combines the
processed information with information about the current needs of the animal and
with memory of past circumstances. Finally, on the basis of the results, it
generates motor response patterns. These signal-processing tasks require intricate
interplay between a variety of functional subsystems.[84]

The function of the brain is to provide coherent control over the actions of an
animal. A centralized brain allows groups of muscles to be co-activated in complex
patterns; it also allows stimuli impinging on one part of the body to evoke
responses in other parts, and it can prevent different parts of the body from
acting at cross-purposes to each other.[84]
Perception
Drawing showing the ear, inner ear, and brain areas involved in hearing. A series
of light blue arrows shows the flow of signals through the system.
Diagram of signal processing in the auditory system

The human brain is provided with information about light, sound, the chemical
composition of the atmosphere, temperature, the position of the body in space
(proprioception), the chemical composition of the bloodstream, and more. In other
animals additional senses are present, such as the infrared heat-sense of snakes,
the magnetic field sense of some birds, or the electric field sense mainly seen in
aquatic animals.

Each sensory system begins with specialized receptor cells,[8] such as

photoreceptor cells in the retina of the eye, or vibration-sensitive hair cells in
the cochlea of the ear. The axons of sensory receptor cells travel into the spinal
cord or brain, where they transmit their signals to a first-order sensory nucleus
dedicated to one specific sensory modality. This primary sensory nucleus sends
information to higher-order sensory areas that are dedicated to the same modality.
Eventually, via a way-station in the thalamus, the signals are sent to the cerebral
cortex, where they are processed to extract the relevant features, and integrated
with signals coming from other sensory systems.[8]
Motor control

Motor systems are areas of the brain that are involved in initiating body
movements, that is, in activating muscles. Except for the muscles that control the
eye, which are driven by nuclei in the midbrain, all the voluntary muscles in the
body are directly innervated by motor neurons in the spinal cord and hindbrain.[8]
Spinal motor neurons are controlled both by neural circuits intrinsic to the spinal
cord, and by inputs that descend from the brain. The intrinsic spinal circuits
implement many reflex responses, and contain pattern generators for rhythmic
movements such as walking or swimming. The descending connections from the brain
allow for more sophisticated control.[8]

The brain contains several motor areas that project directly to the spinal cord. At
the lowest level are motor areas in the medulla and pons, which control stereotyped
movements such as walking, breathing, or swallowing. At a higher level are areas in
the midbrain, such as the red nucleus, which is responsible for coordinating
movements of the arms and legs. At a higher level yet is the primary motor cortex,
a strip of tissue located at the posterior edge of the frontal lobe. The primary
motor cortex sends projections to the subcortical motor areas, but also sends a
massive projection directly to the spinal cord, through the pyramidal tract. This
direct corticospinal projection allows for precise voluntary control of the fine
details of movements. Other motor-related brain areas exert secondary effects by
projecting to the primary motor areas. Among the most important secondary areas are
the premotor cortex, supplementary motor area, basal ganglia, and cerebellum.[8] In
addition to all of the above, the brain and spinal cord contain extensive circuitry
to control the autonomic nervous system which controls the movement of the smooth
muscle of the body.[8]
Major areas involved in controlling movement Area Location Function
Ventral horn Spinal cord Contains motor neurons that directly activate
muscles[85]
Oculomotor nuclei Midbrain Contains motor neurons that directly activate
the eye muscles[86]
Cerebellum Hindbrain Calibrates precision and timing of movements[8]
Basal ganglia Forebrain Action selection on the basis of motivation[87]
Motor cortex Frontal lobe Direct cortical activation of spinal motor
circuits[88]
Premotor cortex Frontal lobe Groups elementary movements into coordinated
patterns[8]
Supplementary motor area Frontal lobe Sequences movements into temporal
patterns[89]
Prefrontal cortex Frontal lobe Planning and other executive
functions[90]
Sleep
Main article: Sleep
See also: Arousal

Many animals alternate between sleeping and waking in a daily cycle. Arousal and
alertness are also modulated on a finer time scale by a network of brain areas.[8]
A key component of the sleep system is the suprachiasmatic nucleus (SCN), a tiny
part of the hypothalamus located directly above the point at which the optic nerves
from the two eyes cross. The SCN contains the body's central biological clock.
Neurons there show activity levels that rise and fall with a period of about 24
hours, circadian rhythms: these activity fluctuations are driven by rhythmic
changes in expression of a set of "clock genes". The SCN continues to keep time
even if it is excised from the brain and placed in a dish of warm nutrient
solution, but it ordinarily receives input from the optic nerves, through the
retinohypothalamic tract (RHT), that allows daily light-dark cycles to calibrate
the clock.[91]

The SCN projects to a set of areas in the hypothalamus, brainstem, and midbrain
that are involved in implementing sleep-wake cycles. An important component of the
system is the reticular formation, a group of neuron-clusters scattered diffusely
through the core of the lower brain. Reticular neurons send signals to the
thalamus, which in turn sends activity-level-controlling signals to every part of
the cortex. Damage to the reticular formation can produce a permanent state of
coma.[8]

Sleep involves great changes in brain activity.[8] Until the 1950s it was generally
believed that the brain essentially shuts off during sleep,[92] but this is now
known to be far from true; activity continues, but patterns become very different.
There are two types of sleep: REM sleep (with dreaming) and NREM (non-REM, usually
without dreaming) sleep, which repeat in slightly varying patterns throughout a
sleep episode. Three broad types of distinct brain activity patterns can be
measured: REM, light NREM and deep NREM. During deep NREM sleep, also called slow
wave sleep, activity in the cortex takes the form of large synchronized waves,
whereas in the waking state it is noisy and desynchronized. Levels of the
neurotransmitters norepinephrine and serotonin drop during slow wave sleep, and
fall almost to zero during REM sleep; levels of acetylcholine show the reverse
pattern.[8]
Homeostasis
Cross-section of a human head, showing location of the hypothalamus

For any animal, survival requires maintaining a variety of parameters of bodily

state within a limited range of variation: these include temperature, water
content, salt concentration in the bloodstream, blood glucose levels, blood oxygen
level, and others.[93] The ability of an animal to regulate the internal
environment of its body—the milieu intérieur, as the pioneering physiologist Claude
Bernard called it—is known as homeostasis (Greek for "standing still").[94]
Maintaining homeostasis is a crucial function of the brain. The basic principle
that underlies homeostasis is negative feedback: any time a parameter diverges from
its set-point, sensors generate an error signal that evokes a response that causes
the parameter to shift back toward its optimum value.[93] (This principle is widely
used in engineering, for example in the control of temperature using a thermostat.)

In vertebrates, the part of the brain that plays the greatest role is the
hypothalamus, a small region at the base of the forebrain whose size does not
reflect its complexity or the importance of its function.[93] The hypothalamus is a
collection of small nuclei, most of which are involved in basic biological
functions. Some of these functions relate to arousal or to social interactions such
as sexuality, aggression, or maternal behaviors; but many of them relate to
homeostasis. Several hypothalamic nuclei receive input from sensors located in the
lining of blood vessels, conveying information about temperature, sodium level,
glucose level, blood oxygen level, and other parameters. These hypothalamic nuclei
send output signals to motor areas that can generate actions to rectify
deficiencies. Some of the outputs also go to the pituitary gland, a tiny gland
attached to the brain directly underneath the hypothalamus. The pituitary gland
secretes hormones into the bloodstream, where they circulate throughout the body
and induce changes in cellular activity.[95]
Motivation
Components of the basal ganglia, shown in two cross-sections of the human brain.
Blue: caudate nucleus and putamen. Green: globus pallidus. Red: subthalamic
nucleus. Black: substantia nigra.

The individual animals need to express survival-promoting behaviors, such as

seeking food, water, shelter, and a mate.[96] The motivational system in the brain
monitors the current state of satisfaction of these goals, and activates behaviors
to meet any needs that arise. The motivational system works largely by a reward–
punishment mechanism. When a particular behavior is followed by favorable
consequences, the reward mechanism in the brain is activated, which induces
structural changes inside the brain that cause the same behavior to be repeated
later, whenever a similar situation arises. Conversely, when a behavior is followed
by unfavorable consequences, the brain's punishment mechanism is activated,
inducing structural changes that cause the behavior to be suppressed when similar
situations arise in the future.[97]

Most organisms studied to date use a reward–punishment mechanism: for instance,

worms and insects can alter their behavior to seek food sources or to avoid
dangers.[98] In vertebrates, the reward-punishment system is implemented by a
specific set of brain structures, at the heart of which lie the basal ganglia, a
set of interconnected areas at the base of the forebrain.[47] The basal ganglia are
the central site at which decisions are made: the basal ganglia exert a sustained
inhibitory control over most of the motor systems in the brain; when this
inhibition is released, a motor system is permitted to execute the action it is
programmed to carry out. Rewards and punishments function by altering the
relationship between the inputs that the basal ganglia receive and the decision-
signals that are emitted. The reward mechanism is better understood than the
punishment mechanism, because its role in drug abuse has caused it to be studied
very intensively. Research has shown that the neurotransmitter dopamine plays a
central role: addictive drugs such as cocaine, amphetamine, and nicotine either
cause dopamine levels to rise or cause the effects of dopamine inside the brain to
be enhanced.[99]
Learning and memory

Almost all animals are capable of modifying their behavior as a result of

experience—even the most primitive types of worms. Because behavior is driven by
brain activity, changes in behavior must somehow correspond to changes inside the
brain. Already in the late 19th century theorists like Santiago Ramón y Cajal
argued that the most plausible explanation is that learning and memory are
expressed as changes in the synaptic connections between neurons.[100] Until 1970,
however, experimental evidence to support the synaptic plasticity hypothesis was
lacking. In 1971 Tim Bliss and Terje Lømo published a paper on a phenomenon now
called long-term potentiation: the paper showed clear evidence of activity-induced
synaptic changes that lasted for at least several days.[101] Since then technical
advances have made these sorts of experiments much easier to carry out, and
thousands of studies have been made that have clarified the mechanism of synaptic
change, and uncovered other types of activity-driven synaptic change in a variety
of brain areas, including the cerebral cortex, hippocampus, basal ganglia, and
cerebellum.[102] Brain-derived neurotrophic factor (BDNF) and physical activity
appear to play a beneficial role in the process.[103]

Neuroscientists currently distinguish several types of learning and memory that are
implemented by the brain in distinct ways:

Working memory is the ability of the brain to maintain a temporary

representation of information about the task that an animal is currently engaged
in. This sort of dynamic memory is thought to be mediated by the formation of cell
assemblies—groups of activated neurons that maintain their activity by constantly
stimulating one another.[104]
Episodic memory is the ability to remember the details of specific events. This
sort of memory can last for a lifetime. Much evidence implicates the hippocampus in
playing a crucial role: people with severe damage to the hippocampus sometimes show
amnesia, that is, inability to form new long-lasting episodic memories.[105]
Semantic memory is the ability to learn facts and relationships. This sort of
memory is probably stored largely in the cerebral cortex, mediated by changes in
connections between cells that represent specific types of information.[106]
Instrumental learning is the ability for rewards and punishments to modify
behavior. It is implemented by a network of brain areas centered on the basal
ganglia.[107]
Motor learning is the ability to refine patterns of body movement by
practicing, or more generally by repetition. A number of brain areas are involved,
including the premotor cortex, basal ganglia, and especially the cerebellum, which
functions as a large memory bank for microadjustments of the parameters of
movement.[108]

Research
Main article: Neuroscience
"Brain research" redirects here. For the scientific journal, see Brain Research.
The Human Brain Project is a large scientific research project, starting in 2013,
which aims to simulate the complete human brain.

The field of neuroscience encompasses all approaches that seek to understand the
brain and the rest of the nervous system.[8] Psychology seeks to understand mind
and behavior, and neurology is the medical discipline that diagnoses and treats
diseases of the nervous system. The brain is also the most important organ studied
in psychiatry, the branch of medicine that works to study, prevent, and treat
mental disorders.[109] Cognitive science seeks to unify neuroscience and psychology
with other fields that concern themselves with the brain, such as computer science
(artificial intelligence and similar fields) and philosophy.[110]

The oldest method of studying the brain is anatomical, and until the middle of the
20th century, much of the progress in neuroscience came from the development of
better cell stains and better microscopes. Neuroanatomists study the large-scale
structure of the brain as well as the microscopic structure of neurons and their
components, especially synapses. Among other tools, they employ a plethora of
stains that reveal neural structure, chemistry, and connectivity. In recent years,
the development of immunostaining techniques has allowed investigation of neurons
that express specific sets of genes. Also, functional neuroanatomy uses medical
imaging techniques to correlate variations in human brain structure with
differences in cognition or behavior.[111]

Neurophysiologists study the chemical, pharmacological, and electrical properties

of the brain: their primary tools are drugs and recording devices. Thousands of
experimentally developed drugs affect the nervous system, some in highly specific
ways. Recordings of brain activity can be made using electrodes, either glued to
the scalp as in EEG studies, or implanted inside the brains of animals for
extracellular recordings, which can detect action potentials generated by
individual neurons.[112] Because the brain does not contain pain receptors, it is
possible using these techniques to record brain activity from animals that are
awake and behaving without causing distress. The same techniques have occasionally
been used to study brain activity in human patients with intractable epilepsy, in
cases where there was a medical necessity to implant electrodes to localize the
brain area responsible for epileptic seizures.[113] Functional imaging techniques
such as fMRI are also used to study brain activity; these techniques have mainly
been used with human subjects, because they require a conscious subject to remain
motionless for long periods of time, but they have the great advantage of being
noninvasive.[114]
Drawing showing a monkey in a restraint chair, a computer monitor, a rototic arm,
and three pieces of computer equipment, with arrows between them to show the flow
of information.
Design of an experiment in which brain activity from a monkey was used to control a
robotic arm[115]

Another approach to brain function is to examine the consequences of damage to

specific brain areas. Even though it is protected by the skull and meninges,
surrounded by cerebrospinal fluid, and isolated from the bloodstream by the blood–
brain barrier, the delicate nature of the brain makes it vulnerable to numerous
diseases and several types of damage. In humans, the effects of strokes and other
types of brain damage have been a key source of information about brain function.
Because there is no ability to experimentally control the nature of the damage,
however, this information is often difficult to interpret. In animal studies, most
commonly involving rats, it is possible to use electrodes or locally injected
chemicals to produce precise patterns of damage and then examine the consequences
for behavior.[116]

Computational neuroscience encompasses two approaches: first, the use of computers

to study the brain; second, the study of how brains perform computation. On one
hand, it is possible to write a computer program to simulate the operation of a
group of neurons by making use of systems of equations that describe their
electrochemical activity; such simulations are known as biologically realistic
neural networks. On the other hand, it is possible to study algorithms for neural
computation by simulating, or mathematically analyzing, the operations of
simplified "units" that have some of the properties of neurons but abstract out
much of their biological complexity. The computational functions of the brain are
studied both by computer scientists and neuroscientists.[117]

Computational neurogenetic modeling is concerned with the study and development of

dynamic neuronal models for modeling brain functions with respect to genes and
dynamic interactions between genes.

Recent years have seen increasing applications of genetic and genomic techniques to
the study of the brain [118] and a focus on the roles of neurotrophic factors and
physical activity in neuroplasticity.[103] The most common subjects are mice,
because of the availability of technical tools. It is now possible with relative
ease to "knock out" or mutate a wide variety of genes, and then examine the effects
on brain function. More sophisticated approaches are also being used: for example,
using Cre-Lox recombination it is possible to activate or deactivate genes in
specific parts of the brain, at specific times.[118]
History
See also: History of neuroscience
Illustration by René Descartes of how the brain implements a reflex response

The oldest brain to have been discovered was in Armenia in the Areni-1 cave
complex. The brain, estimated to be over 5,000 years old, was found in the skull of
a 12 to 14-year-old girl. Although the brains were shriveled, they were well
preserved due to the climate found inside the cave.[119]

Early philosophers were divided as to whether the seat of the soul lies in the
brain or heart. Aristotle favored the heart, and thought that the function of the
brain was merely to cool the blood. Democritus, the inventor of the atomic theory
of matter, argued for a three-part soul, with intellect in the head, emotion in the
heart, and lust near the liver.[120] The unknown author of On the Sacred Disease, a
medical treatise in the Hippocratic Corpus, came down unequivocally in favor of the
brain, writing:
 Men ought to know that from nothing else but the brain come joys, delights,
laughter and sports, and sorrows, griefs, despondency, and lamentations. ... And by
the same organ we become mad and delirious, and fears and terrors assail us, some
by night, and some by day, and dreams and untimely wanderings, and cares that are
not suitable, and ignorance of present circumstances, desuetude, and
unskillfulness. All these things we endure from the brain, when it is not
healthy...
— On the Sacred Disease, attributed to Hippocrates[121]

Andreas Vesalius' Fabrica, published in 1543, showing the base of the human brain,
including optic chiasma, cerebellum, olfactory bulbs, etc.

The Roman physician Galen also argued for the importance of the brain, and
theorized in some depth about how it might work. Galen traced out the anatomical
relationships among brain, nerves, and muscles, demonstrating that all muscles in
the body are connected to the brain through a branching network of nerves. He
postulated that nerves activate muscles mechanically by carrying a mysterious
substance he called pneumata psychikon, usually translated as "animal spirits".
[120] Galen's ideas were widely known during the Middle Ages, but not much further
progress came until the Renaissance, when detailed anatomical study resumed,
combined with the theoretical speculations of René Descartes and those who followed
him. Descartes, like Galen, thought of the nervous system in hydraulic terms. He
believed that the highest cognitive functions are carried out by a non-physical res
cogitans, but that the majority of behaviors of humans, and all behaviors of
animals, could be explained mechanistically.[120]

The first real progress toward a modern understanding of nervous function, though,
came from the investigations of Luigi Galvani (1737–1798), who discovered that a
shock of static electricity applied to an exposed nerve of a dead frog could cause
its leg to contract. Since that time, each major advance in understanding has
followed more or less directly from the development of a new technique of
investigation. Until the early years of the 20th century, the most important
advances were derived from new methods for staining cells.[122] Particularly
critical was the invention of the Golgi stain, which (when correctly used) stains
only a small fraction of neurons, but stains them in their entirety, including cell
body, dendrites, and axon. Without such a stain, brain tissue under a microscope
appears as an impenetrable tangle of protoplasmic fibers, in which it is impossible
to determine any structure. In the hands of Camillo Golgi, and especially of the
Spanish neuroanatomist Santiago Ramón y Cajal, the new stain revealed hundreds of
distinct types of neurons, each with its own unique dendritic structure and pattern
of connectivity.[123]
A drawing on yellowing paper with an archiving stamp in the corner. A spidery tree
branch structure connects to the top of a mass. A few narrow processes follow away
from the bottom of the mass.
Drawing by Santiago Ramón y Cajal of two types of Golgi-stained neurons from the
cerebellum of a pigeon

In the first half of the 20th century, advances in electronics enabled

investigation of the electrical properties of nerve cells, culminating in work by
Alan Hodgkin, Andrew Huxley, and others on the biophysics of the action potential,
and the work of Bernard Katz and others on the electrochemistry of the synapse.
[124] These studies complemented the anatomical picture with a conception of the
brain as a dynamic entity. Reflecting the new understanding, in 1942 Charles
Sherrington visualized the workings of the brain waking from sleep:

The great topmost sheet of the mass, that where hardly a light had twinkled or
moved, becomes now a sparkling field of rhythmic flashing points with trains of
traveling sparks hurrying hither and thither. The brain is waking and with it the
mind is returning. It is as if the Milky Way entered upon some cosmic dance.
Swiftly the head mass becomes an enchanted loom where millions of flashing shuttles
weave a dissolving pattern, always a meaningful pattern though never an abiding
one; a shifting harmony of subpatterns.
— Sherrington, 1942, Man on his Nature[125]

The invention of electronic computers in the 1940s, along with the development of
mathematical information theory, led to a realization that brains can potentially
be understood as information processing systems. This concept formed the basis of
the field of cybernetics, and eventually gave rise to the field now known as
computational neuroscience.[126] The earliest attempts at cybernetics were somewhat
crude in that they treated the brain as essentially a digital computer in disguise,
as for example in John von Neumann's 1958 book, The Computer and the Brain.[127]
Over the years, though, accumulating information about the electrical responses of
brain cells recorded from behaving animals has steadily moved theoretical concepts
in the direction of increasing realism.[126]

One of the most influential early contributions was a 1959 paper titled What the
frog's eye tells the frog's brain: the paper examined the visual responses of
neurons in the retina and optic tectum of frogs, and came to the conclusion that
some neurons in the tectum of the frog are wired to combine elementary responses in
a way that makes them function as "bug perceivers".[128] A few years later David
Hubel and Torsten Wiesel discovered cells in the primary visual cortex of monkeys
that become active when sharp edges move across specific points in the field of
view—a discovery for which they won a Nobel Prize.[129] Follow-up studies in
higher-order visual areas found cells that detect binocular disparity, color,
movement, and aspects of shape, with areas located at increasing distances from the
primary visual cortex showing increasingly complex responses.[130] Other
investigations of brain areas unrelated to vision have revealed cells with a wide
variety of response correlates, some related to memory, some to abstract types of
cognition such as space.[131]

Theorists have worked to understand these response patterns by constructing

mathematical models of neurons and neural networks, which can be simulated using
computers.[126] Some useful models are abstract, focusing on the conceptual
structure of neural algorithms rather than the details of how they are implemented
in the brain; other models attempt to incorporate data about the biophysical
properties of real neurons.[132] No model on any level is yet considered to be a
fully valid description of brain function, though. The essential difficulty is that
sophisticated computation by neural networks requires distributed processing in
which hundreds or thousands of neurons work cooperatively—current methods of brain
activity recording are only capable of isolating action potentials from a few dozen
neurons at a time.[133]

Furthermore, even single neurons appear to be complex and capable of performing

computations.[134] So, brain models that do not reflect this are too abstract to be
representative of brain operation; models that do try to capture this are very
computationally expensive and arguably intractable with present computational
resources. However, the Human Brain Project is trying to build a realistic,
detailed computational model of the entire human brain. The wisdom of this approach
has been publicly contested, with high-profile scientists on both sides of the
argument.

In the second half of the 20th century, developments in chemistry, electron

microscopy, genetics, computer science, functional brain imaging, and other fields
progressively opened new windows into brain structure and function. In the United
States, the 1990s were officially designated as the "Decade of the Brain" to
commemorate advances made in brain research, and to promote funding for such
research.[135]
In the 21st century, these trends have continued, and several new approaches have
come into prominence, including multielectrode recording, which allows the activity
of many brain cells to be recorded all at the same time;[136] genetic engineering,
which allows molecular components of the brain to be altered experimentally;[118]
genomics, which allows variations in brain structure to be correlated with
variations in DNA properties and neuroimaging.[137]
Society and culture
As food
Main article: Brain as food
Gulai otak, beef brain curry from Indonesia

Animal brains are used as food in numerous cuisines.

In rituals

Some archaeological evidence suggests that the mourning rituals of European

Neanderthals also involved the consumption of the brain.[138]

The Fore people of Papua New Guinea are known to eat human brains. In funerary
rituals, those close to the dead would eat the brain of the deceased to create a
sense of immortality. A prion disease called kuru has been traced to this.[139]
See also

Philosophy portal

Brain–computer interface
Central nervous system disease
List of neuroscience databases
Neurological disorder
Optogenetics
Outline of neuroscience
Aging brain

References

Saladin, Kenneth (2011). Human anatomy (3rd ed.). McGraw-Hill. p. 416. ISBN 978-0-
07-122207-5.
von Bartheld, CS; Bahney, J; Herculano-Houzel, S (15 December 2016). "The search
for true numbers of neurons and glial cells in the human brain: A review of 150
years of cell counting". The Journal of Comparative Neurology. 524 (18): 3865–3895.
doi:10.1002/cne.24040. PMC 5063692. PMID 27187682.
Yuste, Rafael; Church, George M. (March 2014). "The new century of the brain"
(PDF). Scientific American. 310 (3): 38–45. Bibcode:2014SciAm.310c..38Y.
doi:10.1038/scientificamerican0314-38. PMID 24660326. Archived from the original
(PDF) on 2014-07-14.
Shepherd, GM (1994). Neurobiology. Oxford University Press. p. 3. ISBN 978-0-19-
508843-4.
Sporns, O (2010). Networks of the Brain. MIT Press. p. 143. ISBN 978-0-262-01469-4.
Başar, E (2010). Brain-Body-Mind in the Nebulous Cartesian System: A Holistic
Approach by Oscillations. Springer. p. 225. ISBN 978-1-4419-6134-1.
Singh, Inderbir (2006). "A Brief Review of the Techniques Used in the Study of
Neuroanatomy". Textbook of Human Neuroanatomy (7th ed.). Jaypee Brothers. p. 24.
ISBN 978-81-8061-808-6.
Kandel, Eric R.; Schwartz, James Harris; Jessell, Thomas M. (2000). Principles of
neural science (4th ed.). New York: McGraw-Hill. ISBN 978-0-8385-7701-1. OCLC
42073108.
Douglas, RJ; Martin, KA (2004). "Neuronal circuits of the neocortex". Annual Review
of Neuroscience. 27: 419–451. doi:10.1146/annurev.neuro.27.070203.144152. PMID
15217339.
Barnett, MW; Larkman, PM (2007). "The action potential". Practical Neurology. 7
(3): 192–197. PMID 17515599.
Shepherd, Gordon M. (2004). "1. Introduction to synaptic circuits". The Synaptic
Organization of the Brain (5th ed.). New York, New York: Oxford University Press
US. ISBN 978-0-19-515956-1.
Williams, RW; Herrup, K (1988). "The control of neuron number". Annual Review of
Neuroscience. 11: 423–453. doi:10.1146/annurev.ne.11.030188.002231. PMID 3284447.
Heisenberg, M (2003). "Mushroom body memoir: from maps to models". Nature Reviews
Neuroscience. 4 (4): 266–275. doi:10.1038/nrn1074. PMID 12671643. S2CID 5038386.
Jacobs, DK; Nakanishi, N; Yuan, D; et al. (2007). "Evolution of sensory structures
in basal metazoa". Integrative and Comparative Biology. 47 (5): 712–723. CiteSeerX
10.1.1.326.2233. doi:10.1093/icb/icm094. PMID 21669752.
Balavoine, G (2003). "The segmented Urbilateria: A testable scenario". Integrative
and Comparative Biology. 43 (1): 137–147. doi:10.1093/icb/43.1.137. PMID 21680418.
Schmidt-Rhaesa, A (2007). The Evolution of Organ Systems. Oxford University Press.
p. 110. ISBN 978-0-19-856669-4.
Kristan, WB Jr.; Calabrese, RL; Friesen, WO (2005). "Neuronal control of leech
behavior". Prog Neurobiol. 76 (5): 279–327. doi:10.1016/j.pneurobio.2005.09.004.
PMID 16260077. S2CID 15773361.
Barnes, RD (1987). Invertebrate Zoology (5th ed.). Saunders College Pub. p. 1. ISBN
978-0-03-008914-5.
Butler, AB (2000). "Chordate Evolution and the Origin of Craniates: An Old Brain in
a New Head". Anatomical Record. 261 (3): 111–125. doi:10.1002/1097-
0185(20000615)261:3<111::AID-AR6>3.0.CO;2-F. PMID 10867629.
Bulloch, TH; Kutch, W (1995). "Are the main grades of brains different principally
in numbers of connections or also in quality?". In Breidbach O (ed.). The nervous
systems of invertebrates: an evolutionary and comparative approach. Birkhäuser. p.
439. ISBN 978-3-7643-5076-5.
"Flybrain: An online atlas and database of the drosophila nervous system". Archived
from the original on 1998-01-09. Retrieved 2011-10-14.
Konopka, RJ; Benzer, S (1971). "Clock Mutants of Drosophila melanogaster". Proc.
Natl. Acad. Sci. U.S.A. 68 (9): 2112–2116. Bibcode:1971PNAS...68.2112K.
doi:10.1073/pnas.68.9.2112. PMC 389363. PMID 5002428.
Shin, Hee-Sup; et al. (1985). "An unusual coding sequence from a Drosophila clock
gene is conserved in vertebrates". Nature. 317 (6036): 445–448.
Bibcode:1985Natur.317..445S. doi:10.1038/317445a0. PMID 2413365. S2CID 4372369.
Heisenberg, M; Heusipp, M; Wanke, C. (1995). "Structural plasticity in the
Drosophila brain". J. Neurosci. 15 (3): 1951–1960. doi:10.1523/JNEUROSCI.15-03-
01951.1995. PMC 6578107. PMID 7891144.
Brenner, Sydney (1974). "The Genetics of CAENORHABDITIS ELEGANS". Genetics. 77 (1):
71–94. doi:10.1093/genetics/77.1.71. PMC 1213120. PMID 4366476.
Hobert, O (2005). The C. elegans Research Community (ed.). "Specification of the
nervous system". WormBook: 1–19. doi:10.1895/wormbook.1.12.1. PMC 4781215. PMID
18050401.
White, JG; Southgate, E; Thomson, JN; Brenner, S (1986). "The Structure of the
Nervous System of the Nematode Caenorhabditis elegans". Philosophical Transactions
of the Royal Society B. 314 (1165): 1–340. Bibcode:1986RSPTB.314....1W.
doi:10.1098/rstb.1986.0056. PMID 22462104.
Jabr, Ferris (2012-10-02). "The Connectome Debate: Is Mapping the Mind of a Worm
Worth It?". Scientific American. Retrieved 2014-01-18.
Hodgkin J (2001). "Caenorhabditis elegans". In Brenner S, Miller JH (eds.).
Encyclopedia of Genetics. Elsevier. pp. 251–256. ISBN 978-0-12-227080-2.
Kandel, ER (2007). In Search of Memory: The Emergence of a New Science of Mind. WW
Norton. pp. 145–150. ISBN 978-0-393-32937-7.
Shu, D.-G.; Conway Morris, S.; Han, J.; Zhang, Z.-F.; Yasui, K.; Janvier, P.; Chen,
L.; Zhang, X.-L.; Liu, J.-N.; et al. (2003). "Head and backbone of the Early
Cambrian vertebrate Haikouichthys". Nature. 421 (6922): 526–529.
Bibcode:2003Natur.421..526S. doi:10.1038/nature01264. PMID 12556891. S2CID 4401274.
Striedter, GF (2005). "Ch. 3: Conservation in vertebrate brains". Principles of
Brain Evolution. Sinauer Associates. ISBN 978-0-87893-820-9.
Armstrong, E (1983). "Relative brain size and metabolism in mammals". Science. 220
(4603): 1302–1304. Bibcode:1983Sci...220.1302A. doi:10.1126/science.6407108. PMID
6407108.
Jerison, Harry J. (1973). Evolution of the Brain and Intelligence. Academic Press.
pp. 55–74. ISBN 978-0-12-385250-2.
Parent, A; Carpenter, MB (1996). "Ch. 1". Carpenter's Human Neuroanatomy. Williams
& Wilkins. ISBN 978-0-683-06752-1.
Pardridge, W (2005). "The Blood-Brain Barrier: Bottleneck in Brain Drug
Development". NeuroRx. 2 (1): 3–14. doi:10.1602/neurorx.2.1.3. PMC 539316. PMID
15717053.
Northcutt, RG (2008). "Forebrain evolution in bony fishes". Brain Research
Bulletin. 75 (2–4): 191–205. doi:10.1016/j.brainresbull.2007.10.058. PMID 18331871.
S2CID 44619179.
Reiner, A; Yamamoto, K; Karten, HJ (2005). "Organization and evolution of the avian
forebrain". The Anatomical Record Part A: Discoveries in Molecular, Cellular, and
Evolutionary Biology. 287 (1): 1080–1102. doi:10.1002/ar.a.20253. PMID 16206213.
Siegel, A; Sapru, HN (2010). Essential Neuroscience. Lippincott Williams & Wilkins.
pp. 184–189. ISBN 978-0-7817-8383-5.
Swaab, Dick F. (2003). The Human Hypothalamus – Basic and Clinical Aspects: Nuclei
of the human hypothalamus. Part I. Elsevier. ISBN 9780444514905. Retrieved 2021-01-
22.
Jones, Edward G. (1985). The Thalamus. University of Michigan: Plenum Press. ISBN
9780306418563.
Knierim, James. "Cerebellum (Section 3, Chapter 5)". Neuroscience Online.
Department of Neurobiology and Anatomy at The University of Texas Health Science
Center at Houston, McGovern Medical School. Archived from the original on 2017-11-
18. Retrieved 22 January 2021.
Saitoh, K; Ménard, A; Grillner, S (2007). "Tectal control of locomotion, steering,
and eye movements in lamprey". Journal of Neurophysiology. 97 (4): 3093–3108.
doi:10.1152/jn.00639.2006. PMID 17303814.
Richard Swann Lull; Harry Burr Ferris; George Howard Parker; James Rowland Angell;
Albert Galloway Keller; Edwin Grant Conklin (1922). The evolution of man: a series
of lectures delivered before the Yale chapter of the Sigma xi during the academic
year 1921–1922. Yale University Press. p. 50.
Puelles, L (2001). "Thoughts on the development, structure and evolution of the
mammalian and avian telencephalic pallium". Philosophical Transactions of the Royal
Society B. 356 (1414): 1583–1598. doi:10.1098/rstb.2001.0973. PMC 1088538. PMID
11604125.
Salas, C; Broglio, C; Rodríguez, F (2003). "Evolution of forebrain and spatial
cognition in vertebrates: conservation across diversity". Brain, Behavior and
Evolution. 62 (2): 72–82. doi:10.1159/000072438. PMID 12937346. S2CID 23055468.
Grillner, S; et al. (2005). "Mechanisms for selection of basic motor programs—roles
for the striatum and pallidum". Trends in Neurosciences. 28 (7): 364–370.
doi:10.1016/j.tins.2005.05.004. PMID 15935487. S2CID 12927634.
Northcutt, RG (1981). "Evolution of the telencephalon in nonmammals". Annual Review
of Neuroscience. 4: 301–350. doi:10.1146/annurev.ne.04.030181.001505. PMID 7013637.
Northcutt, RG (2002). "Understanding vertebrate brain evolution". Integrative and
Comparative Biology. 42 (4): 743–756. doi:10.1093/icb/42.4.743. PMID 21708771.
Barton, RA; Harvey, PH (2000). "Mosaic evolution of brain structure in mammals".
Nature. 405 (6790): 1055–1058. Bibcode:2000Natur.405.1055B. doi:10.1038/35016580.
PMID 10890446. S2CID 52854758.
Aboitiz, F; Morales, D; Montiel, J (2003). "The evolutionary origin of the
mammalian isocortex: Towards an integrated developmental and functional approach".
Behavioral and Brain Sciences. 26 (5): 535–552. doi:10.1017/S0140525X03000128. PMID
15179935. S2CID 6599761.
Romer, AS; Parsons, TS (1977). The Vertebrate Body. Holt-Saunders International. p.
531. ISBN 978-0-03-910284-5.
Roth, G; Dicke, U (2005). "Evolution of the brain and Intelligence". Trends in
Cognitive Sciences. 9 (5): 250–257. doi:10.1016/j.tics.2005.03.005. PMID 15866152.
S2CID 14758763.
Marino, Lori (2004). "Cetacean Brain Evolution: Multiplication Generates
Complexity" (PDF). International Society for Comparative Psychology (17): 1–16.
Archived from the original (PDF) on 2018-09-16. Retrieved 2010-08-29.
Shoshani, J; Kupsky, WJ; Marchant, GH (2006). "Elephant brain Part I: Gross
morphology, functions, comparative anatomy, and evolution". Brain Research
Bulletin. 70 (2): 124–157. doi:10.1016/j.brainresbull.2006.03.016. PMID 16782503.
S2CID 14339772.
Finlay, BL; Darlington, RB; Nicastro, N (2001). "Developmental structure in brain
evolution". Behavioral and Brain Sciences. 24 (2): 263–308.
doi:10.1017/S0140525X01003958. PMID 11530543. S2CID 20978251.
Calvin, William H. (1996). How Brains Think (1st ed.). New York, NY: BasicBooks.
ISBN 978-0-465-07278-1.
Sereno, MI; Dale, AM; Reppas, AM; Kwong, KK; Belliveau, JW; Brady, TJ; Rosen, BR;
Tootell, RBH (1995). "Borders of multiple visual areas in human revealed by
functional magnetic resonance imaging" (PDF). Science. 268 (5212): 889–893.
Bibcode:1995Sci...268..889S. doi:10.1126/science.7754376. PMID 7754376. Archived
(PDF) from the original on 2006-05-23.
Fuster, Joaquín M. (2008). The Prefrontal Cortex (4th ed.). Elsevier. pp. 1–7. ISBN
978-0-12-373644-4.
Purves, Dale.; Lichtman, Jeff W. (1985). Principles of neural development.
Sunderland, Mass.: Sinauer Associates. ISBN 978-0-87893-744-8. OCLC 10798963.
Wong, RO (1999). "Retinal waves and visual system development". Annual Review of
Neuroscience. St. Louis, MO. 22: 29–47. doi:10.1146/annurev.neuro.22.1.29. PMID
10202531.
Rakic, Pasko (2002). "Adult neurogenesis in mammals: an identity crisis". Journal
of Neuroscience. 22 (3): 614–618. doi:10.1523/JNEUROSCI.22-03-00614.2002. PMC
6758501. PMID 11826088.
Ridley, Matt (2004). Nature via Nurture: Genes, Experience, and What Makes Us
Human. HarperCollins. pp. 1–6. ISBN 978-0-06-000678-5.
Wiesel, T (1982). "Postnatal development of the visual cortex and the influence of
environment" (PDF). Nature. 299 (5884): 583–591. Bibcode:1982Natur.299..583W.
CiteSeerX 10.1.1.547.7497. doi:10.1038/299583a0. PMID 6811951. S2CID 38776857.
Archived (PDF) from the original on 2022-10-09.
van Praag, H; Kempermann, G; Gage, FH (2000). "Neural consequences of environmental
enrichment". Nature Reviews Neuroscience. 1 (3): 191–198. doi:10.1038/35044558.
PMID 11257907. S2CID 9750498.
Cooper, JR; Bloom, FE; Roth, RH (2003). The Biochemical Basis of Neuropharmacology.
Oxford University Press US. ISBN 978-0-19-514008-8.
McGeer, PL; McGeer, EG (1989). "Chapter 15, Amino acid neurotransmitters". In G.
Siegel; et al. (eds.). Basic Neurochemistry. University of Michigan: Raven Press.
pp. 311–332. ISBN 978-0-88167-343-2.
Foster, AC; Kemp, JA (2006). "Glutamate- and GABA-based CNS therapeutics". Current
Opinion in Pharmacology. 6 (1): 7–17. doi:10.1016/j.coph.2005.11.005. PMID
16377242.
Frazer, A; Hensler, JG (1999). "Understanding the neuroanatomical organization of
serotonergic cells in the brain provides insight into the functions of this
neurotransmitter". In Siegel, GJ (ed.). Basic Neurochemistry (Sixth ed.).
Lippincott Williams & Wilkins. ISBN 978-0-397-51820-3.
Mehler, MF; Purpura, DP (2009). "Autism, fever, epigenetics and the locus
coeruleus". Brain Research Reviews. 59 (2): 388–392.
doi:10.1016/j.brainresrev.2008.11.001. PMC 2668953. PMID 19059284.
Rang, HP (2003). Pharmacology. Churchill Livingstone. pp. 476–483. ISBN 978-0-443-
07145-4.
Speckmann E, Elger CE (2004). "Introduction to the neurophysiological basis of the
EEG and DC potentials". In Niedermeyer E, Lopes da Silva FH (eds.).
Electroencephalography: Basic Principles, Clinical Applications, and Related
Fields. Lippincott Williams & Wilkins. pp. 17–31. ISBN 978-0-7817-5126-1.
Buzsáki, Gyorgy (2006). Rhythms of the Brain. Oxford University Press. ISBN
9780199828234.
Nieuwenhuys, R; Donkelaar, HJ; Nicholson, C (1998). The Central Nervous System of
Vertebrates, Volume 1. Springer. pp. 11–14. ISBN 978-3-540-56013-5.
Safi, K; Seid, MA; Dechmann, DK (2005). "Bigger is not always better: when brains
get smaller". Biology Letters. 1 (3): 283–286. doi:10.1098/rsbl.2005.0333. PMC
1617168. PMID 17148188.
Mink, JW; Blumenschine, RJ; Adams, DB (1981). "Ratio of central nervous system to
body metabolism in vertebrates: its constancy and functional basis". American
Journal of Physiology (Submitted manuscript). 241 (3): R203–212.
doi:10.1152/ajpregu.1981.241.3.R203. PMID 7282965. Archived from the original on
2020-08-17. Retrieved 2021-02-10.
Raichle, M; Gusnard, DA (2002). "Appraising the brain's energy budget". Proc. Natl.
Acad. Sci. U.S.A. 99 (16): 10237–10239. Bibcode:2002PNAS...9910237R.
doi:10.1073/pnas.172399499. PMC 124895. PMID 12149485.
Mehagnoul-Schipper, DJ; Van Der Kallen, BF; Colier, WNJM; Van Der Sluijs, MC; Van
Erning, LJ; Thijssen, HO; Oeseburg, B; Hoefnagels, WH; Jansen, RW (2002).
"Simultaneous measurements of cerebral oxygenation changes during brain activation
by near-infrared spectroscopy and functional magnetic resonance imaging in healthy
young and elderly subjects". Hum Brain Mapp. 16 (1): 14–23. doi:10.1002/hbm.10026.
PMC 6871837. PMID 11870923.
Ebert, D.; Haller, RG.; Walton, ME. (Jul 2003). "Energy contribution of octanoate
to intact rat brain metabolism measured by 13C nuclear magnetic resonance
spectroscopy". J Neurosci. 23 (13): 5928–5935. doi:10.1523/JNEUROSCI.23-13-
05928.2003. PMC 6741266. PMID 12843297.
Marin-Valencia, I.; Good, LB.; Ma, Q.; Malloy, CR.; Pascual, JM. (Feb 2013).
"Heptanoate as a neural fuel: energetic and neurotransmitter precursors in normal
and glucose transporter I-deficient (G1D) brain". J Cereb Blood Flow Metab. 33 (2):
175–182. doi:10.1038/jcbfm.2012.151. PMC 3564188. PMID 23072752.
Boumezbeur, F.; Petersen, KF.; Cline, GW.; Mason, GF.; Behar, KL.; Shulman, GI.;
Rothman, DL. (Oct 2010). "The contribution of blood lactate to brain energy
metabolism in humans measured by dynamic 13C nuclear magnetic resonance
spectroscopy". J Neurosci. 30 (42): 13983–13991. doi:10.1523/JNEUROSCI.2040-
10.2010. PMC 2996729. PMID 20962220.
Deelchand, DK.; Shestov, AA.; Koski, DM.; Uğurbil, K.; Henry, PG. (May 2009).
"Acetate transport and utilization in the rat brain". J Neurochem. 109 (Suppl 1):
46–54. doi:10.1111/j.1471-4159.2009.05895.x. PMC 2722917. PMID 19393008.
Soengas, JL; Aldegunde, M (2002). "Energy metabolism of fish brain". Comparative
Biochemistry and Physiology B. 131 (3): 271–296. doi:10.1016/S1096-4959(02)00022-2.
PMID 11959012.
Carew, TJ (2000). "Ch. 1". Behavioral Neurobiology: the Cellular Organization of
Natural Behavior. Sinauer Associates. ISBN 978-0-87893-092-0.
Dafny, N. "Anatomy of the spinal cord". Neuroscience Online. Archived from the
original on 2011-10-08. Retrieved 2011-10-10.
Dragoi, V. "Ocular motor system". Neuroscience Online. Archived from the original
on 2011-11-17. Retrieved 2011-10-10.
Gurney, K; Prescott, TJ; Wickens, JR; Redgrave, P (2004). "Computational models of
the basal ganglia: from robots to membranes". Trends in Neurosciences. 27 (8): 453–
459. doi:10.1016/j.tins.2004.06.003. PMID 15271492. S2CID 2148363.
Knierim, James. "Motor Cortex (Section 3, Chapter 3)". Neuroscience Online.
Department of Neurobiology and Anatomy at The University of Texas Health Science
Center at Houston, McGovern Medical School. Retrieved 2021-01-23.
Shima, K; Tanji, J (1998). "Both supplementary and presupplementary motor areas are
crucial for the temporal organization of multiple movements". Journal of
Neurophysiology. 80 (6): 3247–3260. doi:10.1152/jn.1998.80.6.3247. PMID 9862919.
Miller, EK; Cohen, JD (2001). "An integrative theory of prefrontal cortex
function". Annual Review of Neuroscience. 24 (1): 167–202.
doi:10.1146/annurev.neuro.24.1.167. PMID 11283309. S2CID 7301474.
Antle, MC; Silver, R (2005). "Orchestrating time: arrangements of the brain
circadian clock" (PDF). Trends in Neurosciences. 28 (3): 145–151.
doi:10.1016/j.tins.2005.01.003. PMID 15749168. S2CID 10618277. Archived from the
original (PDF) on 2008-10-31.
Kleitman, Nathaniel (1939). Sleep and Wakefulness. Revised and enlarged edition
1963, Reprint edition 1987. Chicago: The University of Chicago Press, Midway
Reprint. ISBN 978-0-226-44073-6.
Dougherty, Patrick. "Hypothalamus: structural organization". Neuroscience Online.
Archived from the original on 2011-11-17. Retrieved 2011-10-11.
Gross, Charles G. (1998). "Claude Bernard and the constancy of the internal
environment" (PDF). The Neuroscientist. 4 (5): 380–385.
doi:10.1177/107385849800400520. S2CID 51424670. Archived from the original (PDF) on
2018-12-08.
Dougherty, Patrick. "Hypothalamic control of pituitary hormone". Neuroscience
Online. Archived from the original on 2011-11-17. Retrieved 2011-10-11.
Chiel, HJ; Beer, RD (1997). "The brain has a body: adaptive behavior emerges from
interactions of nervous system, body, and environment". Trends in Neurosciences. 20
(12): 553–557. doi:10.1016/S0166-2236(97)01149-1. PMID 9416664. S2CID 5634365.
Berridge, KC (2004). "Motivation concepts in behavioral neuroscience". Physiology &
Behavior. 81 (2): 179–209. doi:10.1016/j.physbeh.2004.02.004. PMID 15159167. S2CID
14149019.
Ardiel, EL; Rankin, CH (2010). "An elegant mind: learning and memory in
Caenorhabditis elegans". Learning and Memory. 17 (4): 191–201.
doi:10.1101/lm.960510. PMID 20335372.
Hyman, SE; Malenka, RC (2001). "Addiction and the brain: the neurobiology of
compulsion and its persistence". Nature Reviews Neuroscience. 2 (10): 695–703.
doi:10.1038/35094560. PMID 11584307. S2CID 3333114.
Ramón y Cajal, S (1894). "The Croonian Lecture: La Fine Structure des Centres
Nerveux". Proceedings of the Royal Society. 55 (331–335): 444–468.
Bibcode:1894RSPS...55..444C. doi:10.1098/rspl.1894.0063.
Lømo, T (2003). "The discovery of long-term potentiation". Philosophical
Transactions of the Royal Society B. 358 (1432): 617–620.
doi:10.1098/rstb.2002.1226. PMC 1693150. PMID 12740104.
Malenka, R; Bear, M (2004). "LTP and LTD: an embarrassment of riches". Neuron. 44
(1): 5–21. doi:10.1016/j.neuron.2004.09.012. PMID 15450156. S2CID 79844.
Bos, I; De Boever, P; Int Panis, L; Meeusen, R (2004). "Physical Activity, Air
Pollution and the Brain". Sports Medicine. 44 (11): 1505–1518. doi:10.1007/s40279-
014-0222-6. PMID 25119155. S2CID 207493297.
Curtis, CE; D'Esposito, M (2003). "Persistent activity in the prefrontal cortex
during working memory". Trends in Cognitive Sciences. 7 (9): 415–423. CiteSeerX
10.1.1.457.9723. doi:10.1016/S1364-6613(03)00197-9. PMID 12963473. S2CID 15763406.
Tulving, E; Markowitsch, HJ (1998). "Episodic and declarative memory: role of the
hippocampus". Hippocampus. 8 (3): 198–204. doi:10.1002/(SICI)1098-
1063(1998)8:3<198::AID-HIPO2>3.0.CO;2-G. PMID 9662134. S2CID 18634842.
Martin, A; Chao, LL (2001). "Semantic memory and the brain: structures and
processes". Current Opinion in Neurobiology. 11 (2): 194–201. doi:10.1016/S0959-
4388(00)00196-3. PMID 11301239. S2CID 3700874.
Balleine, BW; Liljeholm, Mimi; Ostlund, SB (2009). "The integrative function of the
basal ganglia in instrumental learning". Behavioural Brain Research. 199 (1): 43–
52. doi:10.1016/j.bbr.2008.10.034. PMID 19027797. S2CID 36521958.
Doya, K (2000). "Complementary roles of basal ganglia and cerebellum in learning
and motor control". Current Opinion in Neurobiology. 10 (6): 732–739.
doi:10.1016/S0959-4388(00)00153-7. PMID 11240282. S2CID 10962570.
Storrow, Hugh A. (1969). Outline of clinical psychiatry. New York: Appleton-
Century-Crofts, Educational Division. ISBN 978-0-390-85075-1. OCLC 47198.
Thagard, Paul (2007). "Cognitive Science". Stanford Encyclopedia of Philosophy
(Revised, 2nd ed.). Retrieved 2021-01-23.
Bear, MF; Connors, BW; Paradiso, MA (2007). "Ch. 2". Neuroscience: Exploring the
Brain. Lippincott Williams & Wilkins. ISBN 978-0-7817-6003-4.
Dowling, JE (2001). Neurons and Networks. Harvard University Press. pp. 15–24. ISBN
978-0-674-00462-7.
Wyllie, E; Gupta, A; Lachhwani, DK (2005). "Ch. 77". The Treatment of Epilepsy:
Principles and Practice. Lippincott Williams & Wilkins. ISBN 978-0-7817-4995-4.
Laureys S, Boly M, Tononi G (2009). "Functional neuroimaging". In Laureys S, Tononi
G (eds.). The Neurology of Consciousness: Cognitive Neuroscience and
Neuropathology. Academic Press. pp. 31–42. ISBN 978-0-12-374168-4.
Carmena, JM; et al. (2003). "Learning to Control a Brain–Machine Interface for
Reaching and Grasping by Primates". PLOS Biology. 1 (2): 193–208.
doi:10.1371/journal.pbio.0000042. PMC 261882. PMID 14624244.
Kolb, B; Whishaw, I (2008). "Ch. 1". Fundamentals of Human Neuropsychology.
Macmillan. ISBN 978-0-7167-9586-5.
Abbott, LF; Dayan, P (2001). "Preface". Theoretical Neuroscience: Computational and
Mathematical Modeling of Neural Systems. MIT Press. ISBN 978-0-262-54185-5.
Tonegawa, S; Nakazawa, K; Wilson, MA (2003). "Genetic neuroscience of mammalian
learning and memory". Philosophical Transactions of the Royal Society B. 358
(1432): 787–795. doi:10.1098/rstb.2002.1243. PMC 1693163. PMID 12740125.
Bower, Bruce (2009-01-12). "Armenian cave yields ancient human brain". ScienceNews.
Retrieved 2021-01-23.
Finger, Stanley (2001). Origins of Neuroscience. Oxford University Press. pp. 14–
15. ISBN 978-0-19-514694-3.
*Hippocrates (2006) [400 BCE], On the Sacred Disease, Translated by Francis Adams,
Internet Classics Archive: The University of Adelaide Library, archived from the
original on September 26, 2007
Bloom FE (1975). Schmidt FO, Worden FG, Swazey JP, Adelman G (eds.). The
Neurosciences, Paths of Discovery. MIT Press. p. 211. ISBN 978-0-262-23072-8.
Shepherd, GM (1991). "Ch.1 : Introduction and Overview". Foundations of the Neuron
Doctrine. Oxford University Press. ISBN 978-0-19-506491-9.
Piccolino, M (2002). "Fifty years of the Hodgkin-Huxley era". Trends in
Neurosciences. 25 (11): 552–553. doi:10.1016/S0166-2236(02)02276-2. PMID 12392928.
S2CID 35465936.
Sherrington, CS (1942). Man on his nature. Cambridge University Press. p. 178. ISBN
978-0-8385-7701-1.
Churchland, PS; Koch, C; Sejnowski, TJ (1993). "What is computational
neuroscience?". In Schwartz EL (ed.). Computational Neuroscience. MIT Press. pp.
46–55. ISBN 978-0-262-69164-2.
von Neumann, J; Churchland, PM; Churchland, PS (2000). The Computer and the Brain.
Yale University Press. pp. xi–xxii. ISBN 978-0-300-08473-3.
Lettvin, JY; Maturana, HR; McCulloch, WS; Pitts, WH (1959). "What the frog's eye
tells the frog's brain" (PDF). Proceedings of the Institute of Radio Engineers. 47
(11): 1940–1951. doi:10.1109/jrproc.1959.287207. S2CID 8739509. Archived from the
original (PDF) on 2011-09-28.
Hubel, DH; Wiesel, TN (2005). Brain and visual perception: the story of a 25-year
collaboration. Oxford University Press US. pp. 657–704. ISBN 978-0-19-517618-6.
Farah, MJ (2000). The Cognitive Neuroscience of Vision. Wiley-Blackwell. pp. 1–29.
ISBN 978-0-631-21403-8.
Engel, AK; Singer, W (2001). "Temporal binding and the neural correlates of sensory
awareness". Trends in Cognitive Sciences. 5 (1): 16–25. doi:10.1016/S1364-
6613(00)01568-0. PMID 11164732. S2CID 11922975.
Dayan, P; Abbott, LF (2005). "Ch.7: Network models". Theoretical Neuroscience. MIT
Press. ISBN 978-0-262-54185-5.
Averbeck, BB; Lee, D (2004). "Coding and transmission of information by neural
ensembles". Trends in Neurosciences. 27 (4): 225–230.
doi:10.1016/j.tins.2004.02.006. PMID 15046882. S2CID 44512482.
Forrest, MD (2014). "Intracellular Calcium Dynamics Permit a Purkinje Neuron Model
to Perform Toggle and Gain Computations Upon its Inputs". Frontiers in
Computational Neuroscience. 8: 86. doi:10.3389/fncom.2014.00086. PMC 4138505. PMID
25191262.
Jones, EG; Mendell, LM (1999). "Assessing the Decade of the Brain". Science. 284
(5415): 739. Bibcode:1999Sci...284..739J. doi:10.1126/science.284.5415.739. PMID
10336393. S2CID 13261978.
Buzsáki, G (2004). "Large-scale recording of neuronal ensembles" (PDF). Nature
Neuroscience. 7 (5): 446–451. doi:10.1038/nn1233. PMID 15114356. S2CID 18538341.
Archived from the original (PDF) on 2006-09-10.
Geschwind, DH; Konopka, G (2009). "Neuroscience in the era of functional genomics
and systems biology". Nature. 461 (7266): 908–915. Bibcode:2009Natur.461..908G.
doi:10.1038/nature08537. PMC 3645852. PMID 19829370.
Connell, Evan S. (2001). The Aztec Treasure House. Counterpoint Press. ISBN 978-1-
58243-162-8.

Collins, S; McLean CA; Masters CL (2001). "Gerstmann-Straussler-Scheinker

syndrome, fatal familial insomnia, and kuru: a review of these less common human
transmissible spongiform encephalopathies". Journal of Clinical Neuroscience. 8
(5): 387–397. doi:10.1054/jocn.2001.0919. PMID 11535002. S2CID 31976428.

External links
Wikiquote has quotations related to Brain.
Wikimedia Commons has media related to Brain.
Wikisource has the text of the 1911 Encyclopædia Britannica article "Brain".

The Brain from Top to Bottom, at McGill University

"The Brain", BBC Radio 4 discussion with Vivian Nutton, Jonathan Sawday &
Marina Wallace (In Our Time, May 8, 2008)
Our Quest to Understand the Brain – with Matthew Cobb Royal Institution
lecture. Archived at Ghostarchive.

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