Neuroscience

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(Redirected from Neurobiology)
For the journal, see Neuroscience (journal).
Drawing by Santiago Ramn y Cajal (1899) of neurons in the pigeon cerebellum
Neuroscience is the scientific study of the nervous system.[1] Traditionally, ne
uroscience is recognized as a branch of biology. However, it is currently an int
erdisciplinary science that collaborates with other fields such as chemistry, co
gnitive science, computer science, engineering, linguistics, mathematics, medici
ne (including neurology), genetics, and allied disciplines including philosophy,
physics, and psychology. It also exerts influence on other fields, such as neur
oeducation,[2] neuroethics, and neurolaw. The term neurobiology is often used in
terchangeably with the term neuroscience, although the former refers specificall
y to the biology of the nervous system, whereas the latter refers to the entire
science of the nervous system, including elements of psychology as well as the p
urely physical sciences.
The scope of neuroscience has broadened to include different approaches used to
study the molecular, cellular, developmental, structural, functional, evolutiona
ry, computational, and medical aspects of the nervous system. The techniques use
d by neuroscientists have also expanded enormously, from molecular and cellular
studies of individual nerve cells to imaging of sensory and motor tasks in the b
rain. Recent theoretical advances in neuroscience have also been aided by the st
udy of neural networks.
As a result of the increasing number of scientists who study the nervous system,
several prominent neuroscience organizations have been formed to provide a foru
m to all neuroscientists and educators. For example, the International Brain Res
earch Organization was founded in 1960,[3] the International Society for Neuroch
emistry in 1963,[4] the European Brain and Behaviour Society in 1968,[5] and the
Society for Neuroscience in 1969.[6]
Contents
1
History
2
Modern neuroscience
2.1
Molecular and cellular neuroscience
2.2
Neural circuits and systems
2.3
Cognitive and behavioral neuroscience
2.3.1 Insufficient study sizes
2.4
Translational research and medicine
3
Major branches
4
Neuroscience organizations
4.1
Public education and outreach
5
See also
6
References
7
Further reading
8
External links
History[edit]
Main article: History of neuroscience
Illustration from Gray's Anatomy (1918) of a lateral view of the human brain, fe
aturing the hippocampus among other neuroanatomical features
The study of the nervous system dates back to ancient Egypt. Evidence of trepana
tion, the surgical practice of either drilling or scraping a hole into the skull
with the purpose of curing headaches or mental disorders or relieving cranial p
ressure, being performed on patients dates back to Neolithic times and has been
found in various cultures throughout the world. Manuscripts dating back to 1700
BC indicated that the Egyptians had some knowledge about symptoms of brain damag

e.[7]
Early views on the function of the brain regarded it to be a "cranial stuffing"
of sorts. In Egypt, from the late Middle Kingdom onwards, the brain was regularl
y removed in preparation for mummification. It was believed at the time that the
heart was the seat of intelligence. According to Herodotus, the first step of m
ummification was to "take a crooked piece of iron, and with it draw out the brai
n through the nostrils, thus getting rid of a portion, while the skull is cleare
d of the rest by rinsing with drugs."[8]
The view that the heart was the source of consciousness was not challenged until
the time of the Greek physician Hippocrates. He believed that the brain was not
only involved with sensation since most specialized organs (e.g., eyes, ears, ton
gue) are located in the head near the brain but was also the seat of intelligence.
Plato also speculated that the brain was the seat of the rational part of the s
oul.[9] Aristotle, however, believed the heart was the center of intelligence an
d that the brain regulated the amount of heat from the heart.[10] This view was
generally accepted until the Roman physician Galen, a follower of Hippocrates an
d physician to Roman gladiators, observed that his patients lost their mental fa
culties when they had sustained damage to their brains.
Abulcasis, Averroes, Avenzoar, and Maimonides, active in the Medieval Muslim wor
ld, described a number of medical problems related to the brain. In Renaissance
Europe, Vesalius (1514 1564), Ren Descartes (1596 1650), and Thomas Willis (1621 1675)
also made several contributions to neuroscience.
The Golgi stain first allowed for the visualization of individual neurons.
In the first half of the 19th century, Jean Pierre Flourens pioneered the experi
mental method of carrying out localized lesions of the brain in living animals d
escribing their effects on motricity, sensibility and behavior. Studies of the b
rain became more sophisticated after the invention of the microscope and the dev
elopment of a staining procedure by Camillo Golgi during the late 1890s. The pro
cedure used a silver chromate salt to reveal the intricate structures of individ
ual neurons. His technique was used by Santiago Ramn y Cajal and led to the forma
tion of the neuron doctrine, the hypothesis that the functional unit of the brai
n is the neuron.[11] Golgi and Ramn y Cajal shared the Nobel Prize in Physiology
or Medicine in 1906 for their extensive observations, descriptions, and categori
zations of neurons throughout the brain. While Luigi Galvani's pioneering work i
n the late 1700s had set the stage for studying the electrical excitability of m
uscles and neurons, it was in the late 19th century that Emil du Bois-Reymond, J
ohannes Peter Mller, and Hermann von Helmholtz demonstrated that the electrical e
xcitation of neurons predictably affected the electrical states of adjacent neur
ons,[citation needed] and Richard Caton found electrical phenomena in the cerebr
al hemispheres of rabbits and monkeys.
In parallel with this research, work with brain-damaged patients by Paul Broca s
uggested that certain regions of the brain were responsible for certain function
s. At the time, Broca's findings were seen as a confirmation of Franz Joseph Gal
l's theory that language was localized and that certain psychological functions
were localized in specific areas of the cerebral cortex.[12][13] The localizatio
n of function hypothesis was supported by observations of epileptic patients con
ducted by John Hughlings Jackson, who correctly inferred the organization of the
motor cortex by watching the progression of seizures through the body. Carl Wer
nicke further developed the theory of the specialization of specific brain struc
tures in language comprehension and production. Modern research through neuroima
ging techniques, still uses the Brodmann cerebral cytoarchitectonic map (referri
ng to study of cell structure) anatomical definitions from this era in continuin
g to show that distinct areas of the cortex are activated in the execution of sp
ecific tasks.[14]

During the 20th century, neuroscience began to be recognized as a distinct acade

mic discipline in its own right, rather than as studies of the nervous system wi
thin other disciplines. Eric Kandel and collaborators have cited David Rioch, Fr
ancis O. Schmitt, and Stephen Kuffler as having played critical roles in establi
shing the field.[15] Rioch originated the integration of basic anatomical and ph
ysiological research with clinical psychiatry at the Walter Reed Army Institute
of Research, starting in the 1950s. During the same period, Schmitt established
a neuroscience research program within the Biology Department at the Massachuset
ts Institute of Technology, bringing together biology, chemistry, physics, and m
athematics. The first freestanding neuroscience department (then called Psychobi
ology) was founded in 1964 at the University of California, Irvine by James L. M
cGaugh.[citation needed] This was followed by the Department of Neurobiology at
Harvard Medical School which was founded in 1966 by Stephen Kuffler.[citation ne
eded]
In 1952, Alan Lloyd Hodgkin and Andrew Huxley presented a mathematical model for
transmission of electrical signals in neurons of the giant axon of a squid, whi
ch they called "action potentials", and how they are initiated and propagated, k
nown as the Hodgkin Huxley model. In 1961 2, Richard FitzHugh and J. Nagumo simplifi
ed Hodgkin Huxley, in what is called the FitzHugh Nagumo model. In 1962, Bernard Kat
z modeled neurotransmission across the space between neurons known as synapses.
Beginning in 1966, Eric Kandel and collaborators examined biochemical changes in
neurons associated with learning and memory storage in Aplysia. In 1981 Catheri
ne Morris and Harold Lecar combined these models in the Morris Lecar model.
Modern neuroscience[edit]
Main articles: Outline of neuroscience and Twentieth century studies in neurosci
ence
Human nervous system
The scientific study of the nervous system has increased significantly during th
e second half of the twentieth century, principally due to advances in molecular
biology, electrophysiology, and computational neuroscience. This has allowed ne
uroscientists to study the nervous system in all its aspects: how it is structur
ed, how it works, how it develops, how it malfunctions, and how it can be change
d. For example, it has become possible to understand, in much detail, the comple
x processes occurring within a single neuron. Neurons are cells specialized for
communication. They are able to communicate with neurons and other cell types th
rough specialized junctions called synapses, at which electrical or electrochemi
cal signals can be transmitted from one cell to another. Many neurons extrude lo
ng thin filaments of protoplasm called axons, which may extend to distant parts
of the body and are capable of rapidly carrying electrical signals, influencing
the activity of other neurons, muscles, or glands at their termination points. A
nervous system emerges from the assemblage of neurons that are connected to eac
h other.
In vertebrates, the nervous system can be split into two parts, the central nerv
ous system (brain and spinal cord), and the peripheral nervous system. In many s
pecies including all vertebrates
the nervous system is the most complex organ sy
stem in the body, with most of the complexity residing in the brain. The human b
rain alone contains around one hundred billion neurons and one hundred trillion
synapses; it consists of thousands of distinguishable substructures, connected t
o each other in synaptic networks whose intricacies have only begun to be unrave
led. The majority of the approximately 20 25,000 genes belonging to the human geno
me are expressed specifically in the brain. Due to the plasticity of the human b
rain, the structure of its synapses and their resulting functions change through
out life.[16] Thus the challenge of making sense of all this complexity is formi
dable.

Molecular and cellular neuroscience[edit]

Main articles: Molecular neuroscience and Cellular neuroscience
Photograph of a stained neuron in a chicken embryo
The study of the nervous system can be done at multiple levels, ranging from the
molecular and cellular levels to the systems and cognitive levels. At the molec
ular level, the basic questions addressed in molecular neuroscience include the
mechanisms by which neurons express and respond to molecular signals and how axo
ns form complex connectivity patterns. At this level, tools from molecular biolo
gy and genetics are used to understand how neurons develop and how genetic chang
es affect biological functions. The morphology, molecular identity, and physiolo
gical characteristics of neurons and how they relate to different types of behav
ior are also of considerable interest.
The fundamental questions addressed in cellular neuroscience include the mechani
sms of how neurons process signals physiologically and electrochemically. These
questions include how signals are processed by neurites
thin extensions from a n
euronal cell body, consisting of dendrites (specialized to receive synaptic inpu
ts from other neurons) and axons (specialized to conduct nerve impulses called a
ction potentials) and somas (the cell bodies of the neurons containing the nucle
us), and how neurotransmitters and electrical signals are used to process inform
ation in a neuron. Another major area of neuroscience is directed at investigati
ons of the development of the nervous system. These questions include the patter
ning and regionalization of the nervous system, neural stem cells, differentiati
on of neurons and glia, neuronal migration, axonal and dendritic development, tr
ophic interactions, and synapse formation.
Computational neurogenetic modeling is concerned with the study and development
of dynamic neuronal models for modeling brain functions with respect to genes an
d dynamic interactions between genes.
Neural circuits and systems[edit]
Main articles: Biological neural network and Systems neuroscience
At the systems level, the questions addressed in systems neuroscience include ho
w neural circuits are formed and used anatomically and physiologically to produc
e functions such as reflexes, multisensory integration, motor coordination, circ
adian rhythms, emotional responses, learning, and memory. In other words, they a
ddress how these neural circuits function and the mechanisms through which behav
iors are generated. For example, systems level analysis addresses questions conc
erning specific sensory and motor modalities: how does vision work? How do songb
irds learn new songs and bats localize with ultrasound? How does the somatosenso
ry system process tactile information? The related fields of neuroethology and n
europsychology address the question of how neural substrates underlie specific a
nimal and human behaviors. Neuroendocrinology and psychoneuroimmunology examine
interactions between the nervous system and the endocrine and immune systems, re
spectively. Despite many advancements, the way networks of neurons produce compl
ex cognitions and behaviors is still poorly understood.
Cognitive and behavioral neuroscience[edit]
Main article: Cognitive neuroscience
At the cognitive level, cognitive neuroscience addresses the questions of how ps
ychological functions are produced by neural circuitry. The emergence of powerfu
l new measurement techniques such as neuroimaging (e.g., fMRI, PET, SPECT), elec
trophysiology, and human genetic analysis combined with sophisticated experiment
al techniques from cognitive psychology allows neuroscientists and psychologists
to address abstract questions such as how human cognition and emotion are mappe
d to specific neural substrates. Although many studies still hold a reductionist
stance looking for the neurobiological basis of cognitive phenomena, recent res
earch shows that there is an interesting interplay between neuroscientific findi
ngs and conceptual research, soliciting and integrating both perspectives. For e

xample, the neuroscience research on empathy solicited an interesting interdisci

plinary debate involving philosophy, psychology and psychopathology.[17] Moreove
r, the neuroscientific identification of multiple memory systems related to diff
erent brain areas has challenged the idea of memory as a literal reproduction of
the past, supporting a view of memory as a generative, constructive and dynamic
process.[18]
Neuroscience is also allied with the social and behavioral sciences as well as n
ascent interdisciplinary fields such as neuroeconomics, decision theory, and soc
ial neuroscience to address complex questions about interactions of the brain wi
th its environment.
Ultimately neuroscientists would like to understand every aspect of the nervous
system, including how it works, how it develops, how it malfunctions, and how it
can be altered or repaired. The specific topics that form the main foci of rese
arch change over time, driven by an ever-expanding base of knowledge and the ava
ilability of increasingly sophisticated technical methods. Over the long term, i
mprovements in technology have been the primary drivers of progress. Development
s in electron microscopy, computers, electronics, functional brain imaging, and
most recently genetics and genomics, have all been major drivers of progress.
Insufficient study sizes[edit]
Most studies in neurology have too few test subjects to be scientifically sure.
Those insufficient size studies are the basis for all domain-specific diagnoses
in neuropsychiatry, since the few large enough studies there are always find ind
ividuals with the brain changes thought to be associated with a mental condition
but without any of the symptoms. The only diagnoses that can be validated throu
gh large enough brain studies are those on serious brain damages and neurodegene
rative diseases that destroy most of the brain.[19][20]
Translational research and medicine[edit]
Further information: Translational research
Parasagittal MRI of the head of a patient with benign familial macrocephaly
Neurology, psychiatry, neurosurgery, psychosurgery, anesthesiology and pain medi
cine, neuropathology, neuroradiology, ophthalmology, otolaryngology, clinical ne
urophysiology, addiction medicine, and sleep medicine are some medical specialti
es that specifically address the diseases of the nervous system. These terms als
o refer to clinical disciplines involving diagnosis and treatment of these disea
ses. Neurology works with diseases of the central and peripheral nervous systems
, such as amyotrophic lateral sclerosis (ALS) and stroke, and their medical trea
tment. Psychiatry focuses on affective, behavioral, cognitive, and perceptual di
sorders. Anesthesiology focuses on perception of pain, and pharmacologic alterat
ion of consciousness. Neuropathology focuses upon the classification and underly
ing pathogenic mechanisms of central and peripheral nervous system and muscle di
seases, with an emphasis on morphologic, microscopic, and chemically observable
alterations. Neurosurgery and psychosurgery work primarily with surgical treatme
nt of diseases of the central and peripheral nervous systems. The boundaries bet
ween these specialties have been blurring recently as they are all influenced by
basic research in neuroscience. Brain imaging also enables objective, biologica
l insights into mental illness, which can lead to faster diagnosis, more accurat
e prognosis, and help assess patient progress over time.[21]
Integrative neuroscience makes connections across these specialized areas of foc
us.
Major branches[edit]
Modern neuroscience education and research activities can be very roughly catego
rized into the following major branches, based on the subject and scale of the s
ystem in examination as well as distinct experimental or curricular approaches.

Individual neuroscientists, however, often work on questions that span several d

istinct subfields.
Branch Description
Affective neuroscience Affective neuroscience is the study of the neural mechan
isms involved in emotion, typically through experimentation on animal models.[22
]
Behavioral neuroscience Behavioral neuroscience (also known as biological psycho
logy, physiological psychology, biopsychology, or psychobiology) is the applicat
ion of the principles of biology (viz., neurobiology) to the study of genetic, p
hysiological, and developmental mechanisms of behavior in humans and non-human a
nimals.
Cellular neuroscience Cellular neuroscience is the study of neurons at a cellu
lar level including morphology and physiological properties.
Clinical neuroscience This consists of medical specialties such as neurology a
nd psychiatry, as well as many allied health professions such as speech-language
pathology. Neurology is the medical specialty that works with disorders of the
nervous system. Psychiatry is the medical specialty that works with the disorder
s of the brain which include various affective, behavioral, cognitive, and percept
ual disorders. (Also see note below.)
Cognitive neuroscience Cognitive neuroscience is the study of the mechanisms un
derlying cognition with a specific focus on the neural substrates of mental proc
esses.
Computational neuroscience
Computational neuroscience is the study of brain
function in terms of the information processing properties of the structures th
at make up the nervous system. Computational neuroscience can also refer to the
use of computer simulations and theoretical models to study the function of the
nervous system.
Cultural neuroscience Cultural neuroscience is the study of how cultural value
s, practices and beliefs shape and are shaped by the mind, brain and genes acros
s multiple timescales.[23]
Developmental neuroscience
Developmental neuroscience studies the processes
that generate, shape, and reshape the nervous system and seeks to describe the
cellular basis of neural development to address underlying mechanisms.
Evolutionary neuroscience
Evolutionary neuroscience is an interdisciplinar
y scientific research field that studies the evolution of nervous systems.
Molecular neuroscience Molecular neuroscience is a branch of neuroscience that
examines the biology of the nervous system with molecular biology, molecular gen
etics, protein chemistry, and related methodologies.
Neuroengineering
Neuroengineering is a discipline within biomedical engin
eering that uses engineering techniques to understand, repair, replace, or enhan
ce neural systems.
Neuroethology Neuroethology is an interdisciplinary branch that studies the ne
ural basis of natural animal behavior.
Neuroheuristics Neuroheuristics (or Neuristics) is a transdisciplinary paradigm
that studies the information processing effected by the brain as an outcome of n
urture versus nature, at the crossing of top-down and bottom-up strategies.
Neuroimaging
Neuroimaging includes the use of various techniques to either di
rectly or indirectly image the structure and function of the brain.
Neuroinformatics
Neuroinformatics is a discipline within bioinformatics t
hat conducts the organization of neuroscience data and application of computatio
nal models and analytical tools.
Neurolinguistics
Neurolinguistics is the study of the neural mechanisms i
n the human brain that control the comprehension, production, and acquisition of
language.
Neurophysics
Neurophysics investigates the fundamentally physical basis for t
he neurons, neural networks and the brain.
Neurophysiology Neurophysiology is the study of the functioning of the nervous s
ystem, generally using physiological techniques that include measurement and sti
mulation with electrodes or optically with ion- or voltage-sensitive dyes or lig

ht-sensitive channels.
Neuropsychology Neuropsychology is a discipline that resides under the umbrellas
of both psychology and neuroscience, and is involved in activities in the arena
s of both basic science and applied science. In psychology, it is most closely a
ssociated with biopsychology, clinical psychology, cognitive psychology, and dev
elopmental psychology. In neuroscience, it is most closely associated with the c
ognitive, behavioral, social, and affective neuroscience areas. In the applied a
nd medical domain, it is related to neurology and psychiatry.
Paleoneurology Paleoneurology is a field which combines techniques used in pale
ontology and archeology to study brain evolution, especially that of the human b
rain.
Social neuroscience
Social neuroscience is an interdisciplinary field devote
d to understanding how biological systems implement social processes and behavio
r, and to using biological concepts and methods to inform and refine theories of
social processes and behavior.
Systems neuroscience
Systems neuroscience is the study of the function of neu
ral circuits and systems.
Neuroscience organizations[edit]
The largest professional neuroscience organization is the Society for Neuroscien
ce (SFN), which is based in the United States but includes many members from oth
er countries. Since its founding in 1969 the SFN has grown steadily: as of 2010
it recorded 40,290 members from 83 different countries.[24] Annual meetings, hel
d each year in a different American city, draw attendance from researchers, post
doctoral fellows, graduate students, and undergraduates, as well as educational
institutions, funding agencies, publishers, and hundreds of businesses that supp
ly products used in research.
Other major organizations devoted to neuroscience include the International Brai
n Research Organization (IBRO), which holds its meetings in a country from a dif
ferent part of the world each year, and the Federation of European Neuroscience
Societies (FENS), which holds a meeting in a different European city every two y
ears. FENS comprises a set of 32 national-level organizations, including the Bri
tish Neuroscience Association, the German Neuroscience Society (Neurowissenschaf
tliche Gesellschaft), and the French Socit des Neurosciences. The first National H
onor Society in Neuroscience, Nu Rho Psi, was founded in 2006.
In 2013, the BRAIN Initiative was announced in the US.
Public education and outreach[edit]
In addition to conducting traditional research in laboratory settings, neuroscie
ntists have also been involved in the promotion of awareness and knowledge about
the nervous system among the general public and government officials. Such prom
otions have been done by both individual neuroscientists and large organizations
. For example, individual neuroscientists have promoted neuroscience education a
mong young students by organizing the International Brain Bee, which is an acade
mic competition for high school or secondary school students worldwide.[25] In t
he United States, large organizations such as the Society for Neuroscience have
promoted neuroscience education by developing a primer called Brain Facts,[26] c
ollaborating with public school teachers to develop Neuroscience Core Concepts f
or K-12 teachers and students,[27] and cosponsoring a campaign with the Dana Fou
ndation called Brain Awareness Week to increase public awareness about the progr
ess and benefits of brain research.[28] In Canada, the CIHR Canadian National Br
ain Bee is held annually at McMaster University.[29]
Finally, neuroscientists have also collaborated with other education experts to
study and refine educational techniques to optimize learning among students, an
emerging field called educational neuroscience.[30] Federal agencies in the Unit
ed States, such as the National Institute of Health (NIH)[31] and National Scien
ce Foundation (NSF),[32] have also funded research that pertains to best practic
es in teaching and learning of neuroscience concepts.

See also[edit]
List of neuroscience databases
List of neuroscience topics
List of neuroscientists
Outline of brain mapping
Outline of the human brain
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Further reading[edit]
Bear, M. F.; B. W. Connors; M. A. Paradiso (2006). Neuroscience: Exploring the B
rain (3rd ed.). Philadelphia: Lippincott. ISBN 0-7817-6003-8.
Binder, Marc D.; Hirokawa, Nobutaka; Windhorst, Uwe, eds. (2009). Encyclopedia o
f Neuroscience. Springer. ISBN 978-3-540-23735-8.
Kandel, ER; Schwartz JH; Jessell TM (2012). Principles of Neural Science (5th ed
.). New York: McGraw-Hill. ISBN 0-8385-7701-6.
Squire, L. et al. (2012). Fundamental Neuroscience, 4th edition. Academic Press;
ISBN 0-12-660303-0
Byrne and Roberts (2004). From Molecules to Networks. Academic Press; ISBN 0-12148660-5
Sanes, Reh, Harris (2005). Development of the Nervous System, 2nd edition. Acade
mic Press; ISBN 0-12-618621-9
Siegel et al. (2005). Basic Neurochemistry, 7th edition. Academic Press; ISBN 012-088397-X
Rieke, F. et al. (1999). Spikes: Exploring the Neural Code. The MIT Press; Repri
nt edition ISBN 0-262-68108-0
section.47 Neuroscience 2nd ed. Dale Purves, George J. Augustine, David Fitzpatr
ick, Lawrence C. Katz, Anthony-Samuel LaMantia, James O. McNamara, S. Mark Willi
ams. Published by Sinauer Associates, Inc., 2001.
section.18 Basic Neurochemistry: Molecular, Cellular, and Medical Aspects 6th ed
. by George J. Siegel, Bernard W. Agranoff, R. Wayne Albers, Stephen K. Fisher,
Michael D. Uhler, editors. Published by Lippincott, Williams & Wilkins, 1999.
Andreasen, Nancy C. (March 4, 2004). Brave New Brain: Conquering Mental Illness
in the Era of the Genome. Oxford University Press. ISBN 978-0-19-514509-0.[dead
link]
Damasio, A. R. (1994). Descartes' Error: Emotion, Reason, and the Human Brain. N
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