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can be said about them in a human context. The most important is brain disease and the eects of brain damage,
covered in the human brain article because the most common diseases of the human brain either do not show up
in other species, or else manifest themselves in dierent
The brain is an organ that serves as the center of the ways.
nervous system in all vertebrate and most invertebrate animals. Only a few invertebrates such as sponges, jellysh,
adult sea squirts and starsh do not have a brain; diuse 1 Anatomy
or localised nerve nets are present instead. The brain is
located in the head, usually close to the primary sensory
organs for such senses as vision, hearing, balance, taste,
and smell. The brain is the most complex organ in a vertebrates body. In a typical human, the cerebral cortex
(the largest part) is estimated to contain 1533 billion
neurons,[1] each connected by synapses to several thousand other neurons. These neurons communicate with
one another by means of long protoplasmic bers called
axons, which carry trains of signal pulses called action
potentials to distant parts of the brain or body targeting
specic recipient cells.
Physiologically, the function of the brain is to exert centralized control over the other organs of the body. The
brain acts 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
reexes 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 shape and size of the brain varies greatly in dierent species, and identifying common features is often
dicult.[3] Nevertheless, there are a number of principles of brain architecture that apply across a wide range
of species.[4] Some aspects of brain structure are common
to almost the entire range of animal species;[5] others disThe operations of individual brain cells are now under- tinguish advanced brains from more primitive ones, or
[3]
stood in considerable detail but the way they cooperate distinguish vertebrates from invertebrates.
[2]
in ensembles of millions is yet to be solved. Recent The simplest way to gain information about brain
models in modern neuroscience treat the brain as a bi- anatomy is by visual inspection, but many more sophistiological computer, very dierent in mechanism from an cated techniques have been developed. Brain tissue in its
electronic computer, but similar in the sense that it ac- natural state is too soft to work with, but it can be hardquires information from the surrounding world, stores it, ened by immersion in alcohol or other xatives, and then
and processes it in a variety of ways, analogous to the sliced apart for examination of the interior. Visually, the
central processing unit (CPU) in a computer.
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 specic 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.[6]
1.1
1 ANATOMY
Cellular structure
Neurons often have extensive networks of dendrites, which receive synaptic connections. Shown is a pyramidal neuron from
the hippocampus, stained for green uorescent protein.
1.2
Evolution
1.2
Evolution
1.2.1
Nerve cord
Anus
Gut
Brain
Mouth
Fruit ies (Drosophila) have been extensively studied to gain insight into the role of genes in brain development.
Invertebrates
This category includes arthropods, molluscs, and numerous types of worms. The diversity of invertebrate
body plans is matched by an equal diversity in brain
structures.[18]
1 ANATOMY
neurons, always in the same places, making identical synaptic connections in every worm.[26] Brenners team sliced worms into thousands of ultrathin
sections and photographed each one under an electron microscope, then visually matched bers 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]
1.2.3
Vertebrates
Mesencephalon
(midbrain)
Metencephalon
Myelencephalon
Rhombencephalon
(hindbrain)
Spinal cord
1.2
Evolution
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 t 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 identied within the vertebrate brain
based on ne distinctions of neural structure, chemistry,
and connectivity.[7]
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 shes (the great majority of existing sh 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 dicult
to match brain components from one species with those
of another species.[38]
5
functions as currently understood:
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.[7]
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 actiongenerating 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.[7] 10% of the brains 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
shes, such as lampreys, this region is the largest part
of the brain.[43] The superior colliculus is part of the
midbrain.
2
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 ssures 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]
DEVELOPMENT
cortex.[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]
7
ucts that inuence axonal pathnding.[60]
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 uid-lled ventricle at the center. At
the front end, the ventricles and cord swell to form three
vesicles that are the precursors of the forebrain, midbrain,
and 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 nal 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 specic 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 pathnding 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 prod-
3 PHYSIOLOGY
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.[7]
3.1
4.1
3.3
Information processing
Metabolism
Functions
10
4 FUNCTIONS
patterns by constructing mathematical models of neurons and neural networks, which can be simulated using computers.[85] 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.[91] No model
on any level is yet considered to be a fully valid description of brain function, though. The essential diculty
is that sophisticated computation by neural networks requires distributed processing in which hundreds or thousands of neurons work cooperativelycurrent methods
of brain activity recording are only capable of isolating
action potentials from a few dozen neurons at a time.[92]
Furthermore, even single neurons appear to be complex
and capable of performing computations.[93] So, brain
models that don't reect 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, having said this, the Human Brain Project is trying
to build a realistic, detailed computational model of the
entire human brain. It remains to be seen what level of
success they can achieve in the time frame of the project
and the wisdom of it has been publicly contested, with
high-prole scientists on both sides of the argument.
4.2
Perception
One of the primary functions of a brain is to extract biologically relevant information from sensory inputs. The
human brain is provided with information about light,
sound, the chemical composition of the atmosphere, temperature, head orientation, limb position, the chemical
composition of the bloodstream, and more. In other
animals additional senses may be present, such as the
infrared heat-sense of snakes, the magnetic eld sense
of some birds, or the electric eld sense of some types
of sh. Moreover, other animals may develop existing
4.5
Homeostasis
11
4.4
Arousal
4.5 Homeostasis
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.[101] The ability of an animal to regulate the internal environment of its body
the milieu intrieur, as pioneering physiologist Claude
Bernard called itis known as homeostasis (Greek for
standing still).[102] 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.[101] (This principle
is widely used in engineering, for example in the control
of temperature using a thermostat.)
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 neuronclusters scattered diusely through the core of the lower
brain. Reticular neurons send signals to the thalamus,
which in turn sends activity-level-controlling signals to In vertebrates, the part of the brain that plays the greatevery part of the cortex. Damage to the reticular forma- est role is the hypothalamus, a small region at the base of
tion can produce a permanent state of coma.[7]
the forebrain whose size does not reect its complexity or
Sleep involves great changes in brain activity.[7] Until the the importance of its function.[101] The hypothalamus is
1950s it was generally believed that the brain essentially a collection of small nuclei, most of which are involved
shuts o during sleep,[100] but this is now known to be in basic biological functions. Some of these functions refar from true; activity continues, but patterns become late to arousal or to social interactions such as sexuality,
very dierent. There are two types of sleep: REM sleep aggression, or maternal behaviors; but many of them re(with dreaming) and NREM (non-REM, usually without late to homeostasis. Several hypothalamic nuclei receive
dreaming) sleep, which repeat in slightly varying patterns input from sensors located in the lining of blood vesthroughout a sleep episode. Three broad types of dis- sels, conveying information about temperature, sodium
tinct brain activity patterns can be measured: REM, light level, glucose level, blood oxygen level, and other paramNREM and deep NREM. During deep NREM sleep, also eters. These hypothalamic nuclei send output signals to
called slow wave sleep, activity in the cortex takes the motor areas that can generate actions to rectify decienform of large synchronized waves, whereas in the waking cies. Some of the outputs also go to the pituitary gland, a
state it is noisy and desynchronized. Levels of the neu- tiny gland attached to the brain directly underneath the
rotransmitters norepinephrine and serotonin drop during hypothalamus. The pituitary gland secretes hormones
slow wave sleep, and fall almost to zero during REM into the bloodstream, where they circulate throughout the
sleep; levels of acetylcholine show the reverse pattern.[7] body and induce changes in cellular activity.[103]
12
4 FUNCTIONS
4.6
Motivation
Almost all animals are capable of modifying their behavior as a result of experienceeven the most primitive types of worms. Because behavior is driven by
brain activity, changes in behavior must somehow correspond to changes inside the brain. Theorists dating
back to Santiago Ramn y Cajal argued that the most
plausible explanation is that learning and memory are expressed as changes in the synaptic connections between
neurons.[108] Until 1970, however, experimental evidence
to support the synaptic plasticity hypothesis was lacking. In 1971 Tim Bliss and Terje Lmo 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.[109]
Since then technical advances have made these sorts of
experiments much easier to carry out, and thousands of
studies have been made that have claried 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.[110] Brain-derived neurotrophic
factor (BDNF) and physical activity appear to play a benecial role in the process.[111]
According to evolutionary theory, individuals are genetically programmed to behave in ways that ensure survival and reproductive success. This overarching goal of
genetic tness translates into a set of specic survivalpromoting behaviors, such as seeking food, water, shelter, and a mate.[104] 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 Neuroscientists currently distinguish several types of
punishment mechanism. When a particular behavior is learning and memory that are implemented by the brain
followed by favorable consequences, the reward mecha- in distinct ways:
nism in the brain is activated, which induces structural
changes inside the brain that cause the same behavior
Working memory is the ability of the brain to
to be repeated later, whenever a similar situation arises.
maintain a temporary representation of information
Conversely, when a behavior is followed by unfavorable
about the task that an animal is currently engaged in.
consequences, the brains punishment mechanism is actiThis sort of dynamic memory is thought to be mevated, inducing structural changes that cause the behavdiated by the formation of cell assembliesgroups
ior to be suppressed when similar situations arise in the
of activated neurons that maintain their activity by
[105]
future.
constantly stimulating one another.[112]
Most organisms studied to date utilize a reward
Episodic memory is the ability to remember the depunishment mechanism: for instance, worms and insects
tails of specic events. This sort of memory can last
can alter their behavior to seek food sources or to avoid
for a lifetime. Much evidence implicates the hipdangers.[106] In vertebrates, the reward-punishment syspocampus in playing a crucial role: people with setem is implemented by a specic set of brain structures, at
vere damage to the hippocampus sometimes show
the heart of which lie the basal ganglia, a set of interconamnesia, that is, inability to form new long-lasting
nected areas at the base of the forebrain.[47] There is subepisodic memories.[113]
stantial evidence that the basal ganglia are the central site
at which decisions are made: the basal ganglia exert a sus Semantic memory is the ability to learn facts and
tained inhibitory control over most of the motor systems
relationships. This sort of memory is probably
in the brain; when this inhibition is released, a motor sysstored largely in the cerebral cortex, mediated by
tem is permitted to execute the action it is programmed
changes in connections between cells that represent
to carry out. Rewards and punishments function by alspecic types of information.[114]
tering the relationship between the inputs that the basal
ganglia receive and the decision-signals that are emitted.
Instrumental learning is the ability for rewards
The reward mechanism is better understood than the punand punishments to modify behavior. It is impleishment mechanism, because its role in drug abuse has
mented by a network of brain areas centered on the
caused it to be studied very intensively. Research has
basal ganglia.[115]
shown that the neurotransmitter dopamine plays a central
Motor learning is the ability to rene patterns of
role: addictive drugs such as cocaine, amphetamine, and
body movement by practicing, or more generally by
nicotine either cause dopamine levels to rise or cause the
repetition. A number of brain areas are involved,
eects of dopamine inside the brain to be enhanced.[107]
13
including the premotor cortex, basal ganglia, and especially the cerebellum, which functions as a large
memory bank for microadjustments of the parameters of movement.[116]
tools are drugs and recording devices. Thousands of experimentally developed drugs aect the nervous system,
some in highly specic 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.[120] Because
5 Research
the brain does not contain pain receptors, it is possible
using these techniques to record brain activity from anMain article: Neuroscience
imals that are awake and behaving without causing disThe eld of neuroscience encompasses all approaches tress. The same techniques have occasionally been used
to study brain activity in human patients suering from
intractable epilepsy, in cases where there was a medical
necessity to implant electrodes to localize the brain area
responsible for epileptic seizures.[121] Functional imaging
techniques such as functional magnetic resonance imaging 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.[122]
Another approach to brain function is to examine the consequences of damage to specic brain areas. Even though
it is protected by the skull and meninges, surrounded by
cerebrospinal uid, and isolated from the bloodstream by
the bloodbrain barrier, the delicate nature of the brain
makes it vulnerable to numerous diseases and several
types of damage. In humans, the eects 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 dicult 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.[124]
Computational neuroscience encompasses two approaches: rst, 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
Neurophysiologists study the chemical, pharmacologi- to simulate the operation of a group of neurons by
cal, and electrical properties of the brain: their primary making use of systems of equations that describe their
14
HISTORY
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.[128] Hippocrates, the father of medicine, came
down unequivocally in favor of the brain. In his treatise
on epilepsy he wrote:
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...
Hippocrates, On the Sacred Disease[129]
History
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.[128] Galens ideas were
widely known during the Middle Ages, but not much further progress came until the Renaissance, when detailed
15
anatomical study resumed, combined with the theoretical speculations of Ren 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.[130]
The rst real progress toward a modern understanding
of nervous function, though, came from the investigations of Luigi Galvani, 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.[131] 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
bers, in which it is impossible to determine any structure. In the hands of Camillo Golgi, and especially of
the Spanish neuroanatomist Santiago Ramn y Cajal, the
new stain revealed hundreds of distinct types of neurons,
each with its own unique dendritic structure and pattern
of connectivity.[132]
7 See also
Braincomputer interface
Central nervous system disease
List of neuroscience databases
Drawing by Santiago Ramn y Cajal of two types of Golgistained neurons from the cerebellum of a pigeon
In the rst 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 ac-
Neurological disorder
Neuroplasticity
Outline of neuroscience
The brain as food
16
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9 Further reading
Brain Valentino Braitenberg,
Scholarpedia,
2(11):2918. doi:10.4249/scholarpedia.2918
21
10
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File:Shark_brain.png Source: https://upload.wikimedia.org/wikipedia/commons/b/b4/Shark_brain.png License: Public domain Contributors: This le is a modied version of an image scanned from p 102 of the book The Soul of Man ", by Paul Carus, Open Court Pub.
Co., 1905. It is in the public domain because the copyright has expired. Original artist: William Skaggs
File:Spike-waves.png Source: https://upload.wikimedia.org/wikipedia/commons/2/26/Spike-waves.png License: CC BY-SA 2.0 Contributors: ? Original artist: ?
File:Vertebrate-brain-regions_small.png Source: https://upload.wikimedia.org/wikipedia/commons/b/be/Vertebrate-brain-regions_
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Vertebrate-brain-regions.png Original artist: Vertebrate-brain-regions.png: Looie496
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