Neuroscience

Neuroscience
Neuroscience is the scientific study of the nervous system

(the brain, spinal cord, and peripheral nervous system), its
functions and disorders.[1][2][3] It is a multidisciplinary
science that combines physiology, anatomy, molecular
biology, developmental biology, cytology, psychology,
physics, computer science, chemistry, medicine, statistics, and
mathematical modeling to understand the fundamental and
emergent properties of neurons, glia and neural
circuits.[4][5][6][7][8] The understanding of the biological basis
of learning, memory, behavior, perception, and consciousness
has been described by Eric Kandel as the "epic challenge" of
the biological sciences.[9]
The scope of neuroscience has broadened over time to

include different approaches used to study the nervous
system at different scales. The techniques used by Drawing by Santiago Ramón y Cajal (1899)
neuroscientists have expanded enormously, from molecular of neurons in the pigeon cerebellum
and cellular studies of individual neurons to imaging of
sensory, motor and cognitive tasks in the brain.
History
The earliest study of the nervous system dates to ancient
Egypt. Trepanation, the surgical practice of either drilling or
scraping a hole into the skull for the purpose of curing head
injuries or mental disorders, or relieving cranial pressure, was
first recorded during the Neolithic period. Manuscripts dating
to 1700 BC indicate that the Egyptians had some knowledge
about symptoms of brain damage.[10]
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 regularly removed in Illustration from Gray's Anatomy (1918) of
a lateral view of the human brain, featuring
preparation for mummification. It was believed at the time that
the hippocampus among other
the heart was the seat of intelligence. According to Herodotus,
neuroanatomical features
the first step of mummification was to "take a crooked piece of
iron, and with it draw out the brain through the nostrils, thus
getting rid of a portion, while the skull is cleared of the rest by rinsing with drugs."[11]
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, tongue) are located in the head near the brain—but was also the seat of
intelligence.[12] Plato also speculated that the brain was the seat of the rational part of the soul.[13] Aristotle,
however, believed the heart was the center of intelligence and that the brain regulated the amount of heat
from the heart.[14] This view was generally accepted until the Roman physician Galen, a follower of
Hippocrates and physician to Roman gladiators, observed that his patients lost their mental faculties when
they had sustained damage to their brains.[15]
Abulcasis, Averroes, Avicenna, Avenzoar, and Maimonides, active in the Medieval Muslim world,
described a number of medical problems related to the brain. In Renaissance Europe, Vesalius (1514–1564),
René Descartes (1596–1650), Thomas Willis (1621–1675) and Jan Swammerdam (1637–1680) also made
several contributions to neuroscience.
Luigi Galvani's pioneering work in the late 1700s set the stage for
studying the electrical excitability of muscles and neurons. In 1843
Emil du Bois-Reymond demonstrated the electrical nature of the
nerve signal,[16] whose speed Hermann von Helmholtz proceeded
to measure,[17] and in 1875 Richard Caton found electrical
phenomena in the cerebral hemispheres of rabbits and monkeys.[18]
Adolf Beck published in 1890 similar observations of spontaneous
electrical activity of the brain of rabbits and dogs.[19] Studies of the
The Golgi stain first allowed for the
brain became more sophisticated after the invention of the visualization of individual neurons.
microscope and the development of a staining procedure by
Camillo Golgi during the late 1890s. The procedure used a silver
chromate salt to reveal the intricate structures of individual neurons. His technique was used by Santiago
Ramón y Cajal and led to the formation of the neuron doctrine, the hypothesis that the functional unit of the
brain is the neuron.[20] Golgi and Ramón y Cajal shared the Nobel Prize in Physiology or Medicine in 1906
for their extensive observations, descriptions, and categorizations of neurons throughout the brain.
In parallel with this research, in 1815 Jean Pierre Flourens induced localized lesions of the brain in living
animals to observe their effects on motricity, sensibility and behavior. Work with brain-damaged patients by
Marc Dax in 1836 and Paul Broca in 1865 suggested that certain regions of the brain were responsible for
certain functions. At the time, these findings were seen as a confirmation of Franz Joseph Gall's theory that
language was localized and that certain psychological functions were localized in specific areas of the
cerebral cortex.[21][22] The localization of function hypothesis was supported by observations of epileptic
patients conducted by John Hughlings Jackson, who correctly inferred the organization of the motor cortex
by watching the progression of seizures through the body. Carl Wernicke further developed the theory of
the specialization of specific brain structures in language comprehension and production. Modern research
through neuroimaging techniques, still uses the Brodmann cerebral cytoarchitectonic map (referring to the
study of cell structure) anatomical definitions from this era in continuing to show that distinct areas of the
cortex are activated in the execution of specific tasks.[23]
During the 20th century, neuroscience began to be recognized as a distinct academic discipline in its own
right, rather than as studies of the nervous system within other disciplines. Eric Kandel and collaborators
have cited David Rioch, Francis O. Schmitt, and Stephen Kuffler as having played critical roles in
establishing the field.[24] Rioch originated the integration of basic anatomical and physiological 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
Massachusetts Institute of Technology, bringing together biology, chemistry, physics, and mathematics. The
first freestanding neuroscience department (then called Psychobiology) was founded in 1964 at the
University of California, Irvine by James L. McGaugh.[25] This was followed by the Department of
Neurobiology at Harvard Medical School, which was founded in 1966 by Stephen Kuffler.[26]
In the process of treating epilepsy, Wilder Penfield

produced maps of the location of various
functions (motor, sensory, memory, vision) in the
brain.[27][28] He summarized his findings in a
1950 book called The Cerebral Cortex of
Man.[29] Wilder Penfield and his co-investigators
Edwin Boldrey and Theodore Rasmussen are
considered to be the originators of the cortical
homunculus.[30]
The understanding of neurons and of nervous

system function became increasingly precise and
molecular during the 20th century. For example,
3-D sensory and motor homunculus models at the
in 1952, Alan Lloyd Hodgkin and Andrew Natural History Museum, London
Huxley presented a mathematical model for the
transmission of electrical signals in neurons of the
giant axon of a squid, which they called "action potentials", and how they are initiated and propagated,
known as the Hodgkin–Huxley model. In 1961–1962, Richard FitzHugh and J. Nagumo simplified
Hodgkin–Huxley, in what is called the FitzHugh–Nagumo model. In 1962, Bernard Katz 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 Catherine Morris and Harold Lecar combined these models in the Morris–Lecar model.
Such increasingly quantitative work gave rise to numerous biological neuron models and models of neural
computation.
As a result of the increasing interest about the nervous system, several prominent neuroscience
organizations have been formed to provide a forum to all neuroscientists during the 20th century. For
example, the International Brain Research Organization was founded in 1961,[31] the International Society
for Neurochemistry in 1963,[32] the European Brain and Behaviour Society in 1968,[33] and the Society for
Neuroscience in 1969.[34] Recently, the application of neuroscience research results has also given rise to
applied disciplines as neuroeconomics,[35] neuroeducation,[36] neuroethics,[37] and neurolaw.[38]
Over time, brain research has gone through philosophical, experimental, and theoretical phases, with work
on neural implants and brain simulation predicted to be important in the future.[39]
Modern neuroscience
The scientific study of the nervous system increased significantly during the second half of the twentieth
century, principally due to advances in molecular biology, electrophysiology, and computational
neuroscience. This has allowed neuroscientists to study the nervous system in all its aspects: how it is
structured, how it works, how it develops, how it malfunctions, and how it can be changed.
For example, it has become possible to understand, in
much detail, the complex processes occurring within a
single neuron. Neurons are cells specialized for
communication. They are able to communicate with
neurons and other cell types through specialized
junctions called synapses, at which electrical or
electrochemical signals can be transmitted from one cell
to another. Many neurons extrude a long thin filament of
axoplasm called an axon, 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 each other in neural circuits, and
networks.
The vertebrate nervous system can be split into two Human nervous system
parts: the central nervous system (defined as the brain
and spinal cord), and the peripheral nervous system. In
many species—including all vertebrates—the nervous system is the most complex organ system in the
body, with most of the complexity residing in the brain. The human brain alone contains around one
hundred billion neurons and one hundred trillion synapses; it consists of thousands of distinguishable
substructures, connected to each other in synaptic networks whose intricacies have only begun to be
unraveled. At least one out of three of the approximately 20,000 genes belonging to the human genome is
expressed mainly in the brain.[40]
Due to the high degree of plasticity of the human brain, the structure of its synapses and their resulting
functions change throughout life.[41]
Making sense of the nervous system's dynamic complexity is a formidable research challenge. 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. Analysis of the nervous system is
therefore performed at multiple levels, ranging from the molecular and cellular levels to the systems and
cognitive levels. The specific topics that form the main focus of research change over time, driven by an
ever-expanding base of knowledge and the availability of increasingly sophisticated technical methods.
Improvements in technology have been the primary drivers of progress. Developments in electron
microscopy, computer science, electronics, functional neuroimaging, and genetics and genomics have all
been major drivers of progress.
Advances in the classification of brain cells have been enabled by electrophysiological recording, single-cell
genetic sequencing, and high-quality microscopy, which have combined into a single method pipeline
called patch-sequencing in which all three methods are simultaneously applied using miniature tools.[42]
The efficiency of this method and the large amounts of data that is generated has allowed researchers to
make some general conclusions about cell types; for example that the human and mouse brain have
different versions of fundamentally the same cell types.[43]
Molecular and cellular neuroscience

Basic questions addressed in molecular neuroscience include the
mechanisms by which neurons express and respond to molecular
signals and how axons form complex connectivity patterns. At this
level, tools from molecular biology and genetics are used to understand
how neurons develop and how genetic changes affect biological
functions.[44] The morphology, molecular identity, and physiological
characteristics of neurons and how they relate to different types of
behavior are also of considerable interest.[45]
Questions addressed in cellular neuroscience include the mechanisms

of how neurons process signals physiologically and electrochemically.
These questions include how signals are processed by neurites and
somas and how neurotransmitters and electrical signals are used to Photograph of a stained neuron in
a chicken embryo
process information in a neuron. Neurites are thin extensions from a
neuronal cell body, consisting of dendrites (specialized to receive
synaptic inputs from other neurons) and axons (specialized to conduct nerve impulses called action
potentials). Somas are the cell bodies of the neurons and contain the nucleus.[46]
Another major area of cellular neuroscience is the investigation of the development of the nervous
system.[47] Questions include the patterning and regionalization of the nervous system, axonal and dendritic
development, trophic interactions, synapse formation and the implication of fractones in neural stem
cells,[48][49] differentiation of neurons and glia (neurogenesis and gliogenesis), and neuronal migration.[50]
Computational neurogenetic modeling is concerned with the development of dynamic neuronal models for
modeling brain functions with respect to genes and dynamic interactions between genes, on the cellular
level (CNGM can also be used to model neural systems as well).[51]
Neural circuits and systems

Systems neuroscience research centers on the structural and
functional architecture of the developing human brain, and the
functions of large-scale brain networks, or functionally-connected
systems within the brain. Alongside brain development, systems
neuroscience also focuses on how the structure and function of the
brain enables or restricts the processing of sensory information,
using learned mental models of the world, to motivate behavior.
Proposed organization of motor-
Questions in systems neuroscience include how neural circuits are semantic neural circuits for action
formed and used anatomically and physiologically to produce language comprehension. Adapted
functions such as reflexes, multisensory integration, motor from Shebani et al. (2013).
coordination, circadian rhythms, emotional responses, learning, and
memory.[52] In other words, this area of research studies how
connections are made and morphed in the brain, and the effect it has on human sensation, movement,
attention, inhibitory control, decision-making, reasoning, memory formation, reward, and emotion
regulation.[53]
Specific areas of interest for the field include observations of how the structure of neural circuits effect skill
acquisition, how specialized regions of the brain develop and change (neuroplasticity), and the development
of brain atlases, or wiring diagrams of individual developing brains.[54]
The related fields of neuroethology and neuropsychology address the question of how neural substrates
underlie specific animal and human behaviors.[55] Neuroendocrinology and psychoneuroimmunology
examine interactions between the nervous system and the endocrine and immune systems, respectively.[56]
Despite many advancements, the way that networks of neurons perform complex cognitive processes and
behaviors is still poorly understood.[57]
Cognitive and behavioral neuroscience

Cognitive neuroscience addresses the questions of how psychological functions are produced by neural
circuitry. The emergence of powerful new measurement techniques such as neuroimaging (e.g., fMRI, PET,
SPECT), EEG, MEG, electrophysiology, optogenetics and human genetic analysis combined with
sophisticated experimental techniques from cognitive psychology allows neuroscientists and psychologists
to address abstract questions such as how cognition and emotion are mapped to specific neural substrates.
Although many studies still hold a reductionist stance looking for the neurobiological basis of cognitive
phenomena, recent research shows that there is an interesting interplay between neuroscientific findings and
conceptual research, soliciting and integrating both perspectives. For example, neuroscience research on
empathy solicited an interesting interdisciplinary debate involving philosophy, psychology and
psychopathology.[58] Moreover, the neuroscientific identification of multiple memory systems related to
different 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.[59]
Neuroscience is also allied with the social and behavioral sciences, as well as with nascent interdisciplinary
fields. Examples of such alliances include neuroeconomics, decision theory, social neuroscience, and
neuromarketing to address complex questions about interactions of the brain with its environment. A study
into consumer responses for example uses EEG to investigate neural correlates associated with narrative
transportation into stories about energy efficiency.[60]
Computational neuroscience
Questions in computational neuroscience can span a wide range of levels of traditional analysis, such as
development, structure, and cognitive functions of the brain. Research in this field utilizes mathematical
models, theoretical analysis, and computer simulation to describe and verify biologically plausible neurons
and nervous systems. For example, biological neuron models are mathematical descriptions of spiking
neurons which can be used to describe both the behavior of single neurons as well as the dynamics of
neural networks. Computational neuroscience is often referred to as theoretical neuroscience.
Nanoparticles in medicine are versatile in treating neurological disorders showing promising results in
mediating drug transport across the blood–brain barrier.[61] Implementing nanoparticles in antiepileptic
drugs enhances their medical efficacy by increasing bioavailability in the bloodstream, as well as offering a
measure of control in release time concentration.[61] Although nanoparticles can assist therapeutic drugs by
adjusting physical properties to achieve desirable effects, inadvertent increases in toxicity often occur in
preliminary drug trials.[62] Furthermore, production of nanomedicine for drug trials is economically
consuming, hindering progress in their implementation. Computational models in nanoneuroscience provide
alternatives to study the efficacy of nanotechnology-based medicines in neurological disorders while
mitigating potential side effects and development costs.[61]
Nanomaterials often operate at length scales between classical and quantum regimes.[63] Due to the
associated uncertainties at the length scales that nanomaterials operate, it is difficult to predict their behavior
prior to in vivo studies.[61] Classically, the physical processes which occur throughout neurons are
analogous to electrical circuits. Designers focus on such analogies and model brain activity as a neural
circuit.[64] Success in computational modeling of neurons have led to the development of stereochemical
models that accurately predict acetylcholine receptor-based synapses operating at microsecond time
scales.[64]
Ultrafine nanoneedles for cellular manipulations are thinner than the smallest single walled carbon
nanotubes. Computational quantum chemistry[65] is used to design ultrafine nanomaterials with highly
symmetrical structures to optimize geometry, reactivity and stability.[63]
Behavior of nanomaterials are dominated by long ranged non-bonding interactions.[66] Electrochemical

processes that occur throughout the brain generate an electric field which can inadvertently affect the
behavior of some nanomaterials.[63] Molecular dynamics simulations can mitigate the development phase of
nanomaterials as well as prevent neural toxicity of nanomaterials following in vivo clinical trials.[62] Testing
nanomaterials using molecular dynamics optimizes nano characteristics for therapeutic purposes by testing
different environment conditions, nanomaterial shape fabrications, nanomaterial surface properties, etc.
without the need for in vivo experimentation.[67] Flexibility in molecular dynamic simulations allows
medical practitioners to personalize treatment. Nanoparticle related data from translational nanoinformatics
links neurological patient specific data to predict treatment response.[66]
Neuroscience and medicine
Clinical neuroscience
Neurology, psychiatry, neurosurgery, psychosurgery, anesthesiology and pain medicine, neuropathology,
neuroradiology, ophthalmology, otolaryngology, clinical neurophysiology, addiction medicine, and sleep
medicine are some medical specialties that specifically address the diseases of the nervous system. These
terms also refer to clinical disciplines involving diagnosis and treatment of these diseases.[68]
Neurology works with diseases of the central and peripheral nervous systems, such as amyotrophic lateral
sclerosis (ALS) and stroke, and their medical treatment. Psychiatry focuses on affective, behavioral,
cognitive, and perceptual disorders. Anesthesiology focuses on perception of pain, and pharmacologic
alteration of consciousness. Neuropathology focuses upon the classification and underlying pathogenic
mechanisms of central and peripheral nervous system and muscle diseases, with an emphasis on
morphologic, microscopic, and chemically observable alterations. Neurosurgery and psychosurgery work
primarily with surgical treatment of diseases of the central and peripheral nervous systems.[69]
Translational research
Recently, the boundaries between various specialties have blurred,
as they are all influenced by basic research in neuroscience. For
example, brain imaging enables objective biological insight into
mental illnesses, which can lead to faster diagnosis, more accurate
prognosis, and improved monitoring of patient progress over
time.[70]
Integrative neuroscience describes the effort to combine models and

information from multiple levels of research to develop a coherent
model of the nervous system. For example, brain imaging coupled
with physiological numerical models and theories of fundamental
mechanisms may shed light on psychiatric disorders.[71] An MRI of a male's head showing
benign familial macrocephaly (head
Another important area of translational research is brain–computer circumference > 60 cm)
interfaces, or machines that are able to communicate and influence
the brain. Brain–computer interfaces (BCIs) are currently being
researched for their potential to repair neural systems and restore certain cognitive functions.[72] However,
some ethical considerations have to be dealt with before they are accepted.[73][74]
Major branches
Modern neuroscience education and research activities can be very roughly categorized into the following
major branches, based on the subject and scale of the system in examination as well as distinct experimental
or curricular approaches. Individual neuroscientists, however, often work on questions that span several
distinct subfields.
List of the major branches of neuroscience
Branch Description
Affective neuroscience is the study of the neural mechanisms involved in emotion,

Affective neuroscience
typically through experimentation on animal models.[75]
Behavioral neuroscience (also known as biological psychology, physiological
psychology, biopsychology, or psychobiology) is the application of the principles of
Behavioral neuroscience biology to the study of genetic, physiological, and developmental mechanisms of
behavior in humans and non-human animals.[76]
Cellular neuroscience is the study of neurons at a cellular level including
Cellular neuroscience
morphology and physiological properties.[77]
The scientific study of the biological mechanisms that underlie the disorders and
Clinical neuroscience
diseases of the nervous system.[78]
Cognitive neuroscience is the study of the biological mechanisms underlying
Cognitive neuroscience
cognition.[78]
Computational
neuroscience Computational neuroscience is the theoretical study of the nervous system.[79]
Cultural neuroscience is the study of how cultural values, practices and beliefs
Cultural neuroscience
shape and are shaped by the mind, brain and genes across multiple timescales.[80]
Developmental neuroscience studies the processes that generate, shape, and
Developmental reshape the nervous system and seeks to describe the cellular basis of neural
neuroscience
development to address underlying mechanisms.[81]
Evolutionary neuroscience Evolutionary neuroscience studies the evolution of nervous systems.[82]

Molecular neuroscience studies the nervous system with molecular biology,
Molecular neuroscience
molecular genetics, protein chemistry, and related methodologies.[83]
Nanoneuroscience An interdisciplinary field that integrates nanotechnology and neuroscience.[84]
Neural engineering uses engineering techniques to interact with, understand, repair,

Neural engineering
replace, or enhance neural systems.[85]
Neuroanatomy Neuroanatomy is the study of the anatomy of nervous systems.[86]

Neurochemistry is the study of how neurochemicals interact and influence the
Neurochemistry
function of neurons.[87]
Neuroethology Neuroethology is the study of the neural basis of non-human animals behavior.
Neurogastronomy is the study of flavor and how it affects sensation, cognition, and
Neurogastronomy
memory.[88]
Neurogenetics is the study of the genetical basis of the development and function
Neurogenetics
of the nervous system.[89]
Neuroimaging includes the use of various techniques to either directly or indirectly
Neuroimaging
image the structure and function of the brain.[90]
Neuroimmunology is concerned with the interactions between the nervous and the
Neuroimmunology
immune system.[91]
Neuroinformatics is a discipline within bioinformatics that conducts the organization
Neuroinformatics of neuroscience data and application of computational models and analytical
tools.[92]
Neurolinguistics is the study of the neural mechanisms in the human brain that
Neurolinguistics
control the comprehension, production, and acquisition of language.[93][78]
Neuro-ophthalmology is an academically oriented subspecialty that merges the
Neuro-ophthalmology fields of neurology and ophthalmology, often dealing with complex systemic
diseases that have manifestations in the visual system.
Neurophysics is the branch of biophysics dealing with the development and use of
Neurophysics
physical methods to gain information about the nervous system.[94]
Neurophysiology is the study of the structure and function of the nervous system,
generally using physiological techniques that include measurement and stimulation
Neurophysiology
with electrodes or optically with ion- or voltage-sensitive dyes or light-sensitive
channels.[95]
Neuropsychology is a discipline that resides under the umbrellas of both
psychology and neuroscience, and is involved in activities in the arenas of both
basic science and applied science. In psychology, it is most closely associated
Neuropsychology with biopsychology, clinical psychology, cognitive psychology, and developmental
psychology. In neuroscience, it is most closely associated with the cognitive,
behavioral, social, and affective neuroscience areas. In the applied and medical
domain, it is related to neurology and psychiatry.[96]
Neuropsychopharmacology is interdisciplinary science related to
Neuropsychopharmacology psychopharmacology and fundamental neuroscience, is the study of the neural
mechanisms that drugs act upon to influence behavior.[97]
Paleoneurobiology is a field that combines techniques used in paleontology and
Paleoneurobiology
archeology to study brain evolution, especially that of the human brain.[98]
Social neuroscience is an interdisciplinary field devoted to understanding how
biological systems implement social processes and behavior, and to using
Social neuroscience biological concepts and methods to inform and refine theories of social processes
and behavior.[99]
Systems neuroscience is the study of the function of neural circuits and
Systems neuroscience
systems.[100]
Careers in Neuroscience [101]
Bachelor's Level
Pharmaceutical Sales Residential Counselor
Laboratory Technician Regulatory Affairs Specialist

Psychometrist* Medical Technician*
Science Writer Clinical Research Assistant
Science Advocacy Special Education Assistant

Nonprofit Work Patient Care Assistant*
Health Educator Orthotic and Prosthetic Technician*
EEG Technologist* Lab Animal Care Technician

Medical and Healthcare Manager Sales Engineer
Forensic Science Technician Law Enforcement
Pharmacy Technician* Natural Sciences Manager

Public Policy Advertising/Marketing
Master's Level
Nurse Practitioner Neuroimaging Technician
Physician's Assistant Teacher
Genetic Counselor Epidemiology

Occupational Therapist Biostatistician
Orthotist/Prosthetist Speech-Language Pathologist
Neural Engineer Public Health
Advanced Degree
Medicine (MD, DO) Food Scientist
Research Scientist Pharmacist
Dentist Veterinarian
Physical Therapist Audiologist
Optometrist Lawyer
Clinical Psychologist Professor

Neuropsychologist Chiropractor
Neuroscience organizations
The largest professional neuroscience organization is the Society for Neuroscience (SFN), which is based in
the United States but includes many members from other countries. Since its founding in 1969 the SFN has
grown steadily: as of 2010 it recorded 40,290 members from 83 countries.[102] Annual meetings, held each
year in a different American city, draw attendance from researchers, postdoctoral fellows, graduate students,
and undergraduates, as well as educational institutions, funding agencies, publishers, and hundreds of
businesses that supply products used in research.
Other major organizations devoted to neuroscience include the International Brain Research Organization
(IBRO), which holds its meetings in a country from a different 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 years. FENS comprises a set of 32 national-level organizations, including the British
Neuroscience Association, the German Neuroscience Society (Neurowissenschaftliche Gesellschaft), and
the French Société des Neurosciences.[103] The first National Honor Society in Neuroscience, Nu Rho Psi,
was founded in 2006. Numerous youth neuroscience societies which support undergraduates, graduates and
early career researchers also exist, such as Simply Neuroscience[104] and Project Encephalon.[105]
In 2013, the BRAIN Initiative was announced in the US. The International Brain Initiative[106] was created
in 2017,[107] currently integrated by more than seven national-level brain research initiatives (US, Europe,
Allen Institute, Japan, China, Australia,[108] Canada,[109] Korea,[110] and Israel[111])[112] spanning four
continents.
Public education and outreach

In addition to conducting traditional research in laboratory settings, neuroscientists have also been involved
in the promotion of awareness and knowledge about the nervous system among the general public and
government officials. Such promotions have been done by both individual neuroscientists and large
organizations. For example, individual neuroscientists have promoted neuroscience education among young
students by organizing the International Brain Bee, which is an academic competition for high school or
secondary school students worldwide.[113] In the United States, large organizations such as the Society for
Neuroscience have promoted neuroscience education by developing a primer called Brain Facts,[114]
collaborating with public school teachers to develop Neuroscience Core Concepts for K-12 teachers and
students,[115] and cosponsoring a campaign with the Dana Foundation called Brain Awareness Week to
increase public awareness about the progress and benefits of brain research.[116] In Canada, the CIHR
Canadian National Brain Bee is held annually at McMaster University.[117]
Neuroscience educators formed Faculty for Undergraduate Neuroscience (FUN) in 1992 to share best
practices and provide travel awards for undergraduates presenting at Society for Neuroscience
meetings.[118]
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.[119]
Federal agencies in the United States, such as the National Institute of Health (NIH)[120] and National
Science Foundation (NSF),[121] have also funded research that pertains to best practices in teaching and
learning of neuroscience concepts.
Engineering applications of neuroscience
Neuromorphic computer chips

Neuromorphic engineering is a branch of neuroscience that deals with creating functional physical models
of neurons for the purposes of useful computation. The emergent computational properties of neuromorphic
computers are fundamentally different from conventional computers in the sense that they are a complex
system, and that the computational components are interrelated with no central processor.[122]
One example of such a computer is the SpiNNaker supercomputer.[123]
Sensors can also be made smart with neuromorphic technology. An example of this is the Event Camera's
BrainScaleS (brain-inspired Multiscale Computation in Neuromorphic Hybrid Systems), a hybrid analog
neuromorphic supercomputer located at Heidelberg University in Germany. It was developed as part of the
Human Brain Project's neuromorphic computing platform and is the complement to the SpiNNaker
supercomputer, which is based on digital technology. The architecture used in BrainScaleS mimics
biological neurons and their connections on a physical level; additionally, since the components are made of
silicon, these model neurons operate on average 864 times (24 hours of real time is 100 seconds in the
machine simulation) that of their biological counterparts.[124]
Recent advances in neuromorphic microchip technology have led a group of scientists to create an artificial
neuron that can replace real neurons in diseases.[125][126]
Nobel prizes related to neuroscience

Year Prize field Image Laureate Lifetime Country Rationale Ref.
"in recognition
of his work on
the physiology
of digestion,
Ivan through which
1849– [127]
1904 Physiology Petrovich Russian Empire knowledge on
1936
Pavlov vital aspects of
the subject has
been
transformed and
enlarged"
Camillo 1843–
Kingdom of Italy
Golgi 1926
"in recognition
of their work on
1906 Physiology the structure of [128]
the nervous
system"
Santiago
1852–
Ramón y Restoration (Spain)
1934
Cajal
"for his work on

Allvar 1862– [129]
1911 Physiology Sweden the dioptrics of
Gullstrand 1930
the eye"
"for his work on

the physiology
Robert 1876– [130]
1914 Physiology Austria-Hungary and pathology of
Bárány 1936
the vestibular
apparatus"
Charles
1857–
Scott United Kingdom
1952
Sherrington
"for their
discoveries
1932 Physiology regarding the [131]
functions of
neurons"
Edgar
1889–
Douglas United Kingdom
1977
Adrian
Henry 1875–
United Kingdom
Hallett Dale 1968
"for their
discoveries
relating to [132]
1936 Physiology
chemical
transmission of
nerve impulses"
1873– Austria
Otto Loewi
1961 Germany
"for the
discovery of the
Corneille role played by
Jean 1892– the sinus and [133]
1938 Physiology Belgium
François 1968 aortic
Heymans mechanisms in
the regulation of
respiration"
Joseph 1874–
United States
Erlanger 1965 "for their
discoveries
relating to the
highly [134]
1944 Physiology
differentiated
functions of
Herbert single nerve
1888– fibres"
Spencer United States
1963
Gasser
"for his
discovery of the
functional
organization of
Walter 1881– [135]
Switzerland the interbrain as
Rudolf Hess 1973
a coordinator of
the activities of
the internal
1949 Physiology organs"
"for his
discovery of the
António therapeutic
1874– [135]
Caetano Portugal value of
1955
Egas Moniz leucotomy in
certain
psychoses"
1955 Chemistry Vincent du 1901– United States "for his work on [136]
Vigneaud 1978 biochemically
important
sulphur
compounds,
especially for
the first
synthesis of a
polypeptide
hormone"
(Oxytocin)
"for his
discoveries
relating to
synthetic
compounds that
inhibit the action
Daniel 1907– [137]
1957 Physiology Italy of certain body
Bovet 1992
substances, and
especially their
action on the
vascular system
and the skeletal
muscles"
"for his
discoveries of
the physical
Georg von 1899– [138]
1961 Physiology United States mechanism of
Békésy 1972
stimulation
within the
cochlea"
John Carew 1903–

Australia
Eccles 1997
"for their
discoveries
concerning the
ionic
mechanisms
Alan Lloyd 1914– involved in
1963 Physiology United Kingdom [139]
Hodgkin 1998 excitation and
inhibition in the
peripheral and
central portions
of the nerve cell
membrane"
Andrew
1917–
Fielding United Kingdom
2012
Huxley
1967 Physiology "for their [140]

discoveries
concerning the
Ragnar 1900– Finland
primary
Granit 1991 Sweden
physiological
and chemical
visual
processes in the
eye"
Haldan
1903–
Keffer United States
1983
Hartline
George 1906–
United States
Wald 1997
Julius 1912–
United States "for their
Axelrod 2004
discoveries
concerning the
humoral
transmittors in
Ulf von 1905– [139]
1970 Physiology Sweden the nerve
Euler 1983
terminals and
the mechanism
for their storage,
release and
Bernard 1911– inactivation"
United Kingdom
Katz 2003
Karl von 1886–

Austria
Frisch 1982
"for their
discoveries
concerning
organization and [141]
1973 Physiology
Konrad 1903– elicitation of
Austria
Lorenz 1989 individual and
social behaviour
patterns"
Nikolaas 1907–
Netherlands
Tinbergen 1988
Roger 1924–
France
Guillemin 2024
"for their
discoveries
concerning the [142]
1977 Physiology
peptide hormone
production of
the brain"
Andrew V.
1926– Poland
Schally
1981 Physiology "for his

discoveries
concerning the
Roger W. 1913– [140]
United States functional
Sperry 1994
specialization of
the cerebral
hemispheres"
David H. 1926–
Canada
Hubel 2013
"for their
discoveries
concerning
information [140]
processing in
the visual
system"
Torsten N.
1924– Sweden
Wiesel
Stanley 1922–
United States
Cohen 2020
"for their
1986 Physiology discoveries of [143]
growth factors"
Rita Levi- 1909–

Italy
Montalcini 2012
"for his
discovery of
Stanley B. Prions - a new [144]
1997 Physiology 1942– United States
Prusiner biological
principle of
infection"
"for the first

discovery of an
Jens C. 1918– ion-transporting [145]
1997 Chemistry Denmark
Skou 2018
enzyme, Na+,
K+ -ATPase"
Arvid 1923–
Sweden
Carlsson 2018
"for their
discoveries
concerning
Paul 1925– [146]
2000 Physiology United States signal
Greengard 2019
transduction in
the nervous
system"
Eric R.
1929– United States
Kandel
2003 Chemistry Roderick 1956– United States "for discoveries [147]

MacKinnon concerning
channels in cell
membranes [...]
for structural
and mechanistic
studies of ion
channels"
Richard
Axel
"for their
discoveries of
odorant
2004 Physiology receptors and [148]
the organization
of the olfactory
Linda B. system"
Buck
Robert
Lefkowitz
"for studies of
G-protein- [149]
2012 Chemistry
coupled
receptors""
Brian
Kobilka
John United States

1939–
O'Keefe United Kingdom
"for their
discoveries of
place and grid
cells that [150]
2014 Physiology May-Britt
1963– Norway constitute a
Moser positioning
system in the
brain"
Edvard I.
1962– Norway
Moser
2017 Physiology "for their [151]

discoveries of
Jeffrey C.
1939– United States molecular
Hall
mechanisms
controlling the
circadian
rhythm"
Michael
Rosbash
Michael W.
Young
David Julius 1955– United States "for their

discoveries of
2021 Physiology receptors for [152]
temperature and
touch"
Ardem
1967– Lebanon
Patapoutian
See also
List of neuroscience databases
List of neuroscience journals
List of neuroscientists
Outline of brain mapping
Outline of the human brain
List of regions in the human brain
Gut–brain axis
Connectomics
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Further reading
Bear, M. F.; B. W. Connors; M. A. Paradiso (2006). Neuroscience: Exploring the Brain (https://
archive.org/details/neuroscienceexpl00mark) (3rd ed.). Philadelphia: Lippincott. ISBN 978-0-
7817-6003-4.Binder, Marc D.; Hirokawa, Nobutaka; Windhorst, Uwe, eds. (2009).
Encyclopedia of Neuroscience (https://www.springer.com/biomed/neuroscience/book/978-3-
540-23735-8). 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 978-0-8385-7701-1.
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 0-12-
148660-5
Sanes, Reh, Harris (2005). Development of the Nervous System, 2nd edition. Academic
Press; ISBN 0-12-618621-9
Siegel et al. (2005). Basic Neurochemistry, 7th edition. Academic Press; ISBN 0-12-088397-
X
Rieke, F. et al. (1999). Spikes: Exploring the Neural Code. The MIT Press; Reprint edition
ISBN 0-262-68108-0
section.47 Neuroscience (https://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Search&db=b
ooks&doptcmdl=GenBookHL&term=The+Cellular+Components+of+the+Nervous+System+
AND+neurosci%5Bbook%5D+AND+231002%5Buid%5D&rid=neurosci.) Archived (https://w
eb.archive.org/web/20221212181526/https://www.ncbi.nlm.nih.gov/books?cmd=Search&do
ptcmdl=GenBookHL&term=The+Cellular+Components+of+the+Nervous+System+AND+neu
rosci%5Bbook%5D+AND+231002%5Buid%5D&rid=neurosci.) 2022-12-12 at the Wayback
Machine 2nd ed. Dale Purves, George J. Augustine, David Fitzpatrick, Lawrence C. Katz,
Anthony-Samuel LaMantia, James O. McNamara, S. Mark Williams. Published by Sinauer
Associates, Inc., 2001.
section.18 Basic Neurochemistry: Molecular, Cellular, and Medical Aspects (https://www.ncb
i.nlm.nih.gov/entrez/query.fcgi?cmd=Search&db=books&doptcmdl=GenBookHL&term=Char
acteristics+of+the+Neuron+AND+bnchm%5Bbook%5D+AND+160014%5Buid%5D&rid=bnc
hm.) Archived (https://web.archive.org/web/20221212181525/https://www.ncbi.nlm.nih.gov/b
ooks?cmd=Search&doptcmdl=GenBookHL&term=Characteristics+of+the+Neuron+AND+bn
chm%5Bbook%5D+AND+160014%5Buid%5D&rid=bnchm.) 2022-12-12 at the Wayback
Machine 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 (https://archive.org/details/bravenewbraincon00andr). Oxford University
Press. ISBN 978-0-19-514509-0.
Damasio, A. R. (1994). Descartes' Error: Emotion, Reason, and the Human Brain. New York,
Avon Books. ISBN 0-399-13894-3 (Hardcover) ISBN 0-380-72647-5 (Paperback)
Gardner, H. (1976). The Shattered Mind: The Person After Brain Damage. New York, Vintage
Books, 1976 ISBN 0-394-71946-8
Goldstein, K. (2000). The Organism. New York, Zone Books. ISBN 0-942299-96-5
(Hardcover) ISBN 0-942299-97-3 (Paperback)
Lauwereyns, Jan (February 2010). The Anatomy of Bias: How Neural Circuits Weigh the
Options (https://mitpress.mit.edu/9780262123105). Cambridge, Massachusetts: The MIT
Press. ISBN 978-0-262-12310-5.
Subhash Kak, The Architecture of Knowledge: Quantum Mechanics, Neuroscience,
Computers and Consciousness, Motilal Banarsidass, 2004, ISBN 81-87586-12-5
Llinas R. (2001). I of the vortex: from neurons to self MIT Press. ISBN 0-262-12233-2
Luria, A. R. (1997). The Man with a Shattered World: The History of a Brain Wound.
Cambridge, Massachusetts, Harvard University Press. ISBN 0-224-00792-0 (Hardcover)
ISBN 0-674-54625-3 (Paperback)
Luria, A. R. (1998). The Mind of a Mnemonist: A Little Book About A Vast Memory. New York,
Basic Books, Inc. ISBN 0-674-57622-5
Medina, J. (2008). Brain Rules: 12 Principles for Surviving and Thriving at Work, Home, and
School. Seattle, Pear Press. ISBN 0-9797777-0-4 (Hardcover with DVD)
Pinker, S. (1999). How the Mind Works. W. W. Norton & Company. ISBN 0-393-31848-6
Pinker, S. (2002). The Blank Slate: The Modern Denial of Human Nature. Viking Adult.
ISBN 0-670-03151-8
Robinson, D. L. (2009). Brain, Mind and Behaviour: A New Perspective on Human Nature
(2nd ed.). Dundalk, Ireland: Pontoon Publications. ISBN 978-0-9561812-0-6.
Penrose, R., Hameroff, S. R., Kak, S., & Tao, L. (2011). Consciousness and the universe:
Quantum physics, evolution, brain & mind. Cambridge, MA: Cosmology Science Publishers.
Ramachandran, V. S. (1998). Phantoms in the Brain. New York, HarperCollins. ISBN 0-688-
15247-3 (Paperback)
Rose, S. (2006). 21st Century Brain: Explaining, Mending & Manipulating the Mind ISBN 0-
09-942977-2 (Paperback)
Sacks, O. The Man Who Mistook His Wife for a Hat. Summit Books ISBN 0-671-55471-9
Sacks, O. (1990). Awakenings. New York, Vintage Books. (See also Oliver Sacks) ISBN 0-
671-64834-9 (Hardcover) ISBN 0-06-097368-4 (Paperback)
Encyclopedia:Neuroscience (http://www.scholarpedia.org/article/Encyclopedia:Neuroscienc
e) Archived (https://web.archive.org/web/20200222104731/http://www.scholarpedia.org/articl
e/Encyclopedia:Neuroscience) 2020-02-22 at the Wayback Machine Scholarpedia Expert
articles
Sternberg, E. (2007) Are You a Machine? The Brain, the Mind and What it Means to be
Human. Amherst, New York: Prometheus Books.
Churchland, P. S. (2011) Braintrust: What Neuroscience Tells Us about Morality (http://www.t
hemontrealreview.com/2009/What-neuroscience-tells-us-about-morality.php) Archived (http
s://web.archive.org/web/20201112034251/http://www.themontrealreview.com/2009/What-ne
uroscience-tells-us-about-morality.php) 2020-11-12 at the Wayback Machine. Princeton
University Press. ISBN 0-691-13703-X
Selvin, Paul (2014). "Hot Topics presentation: New Small Quantum Dots for Neuroscience"
(http://spie.org/x106518.xml). SPIE Newsroom. doi:10.1117/2.3201403.17 (https://doi.org/10.
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External links
Neuroscience (https://www.bbc.co.uk/programmes/b00fbd26) on In Our Time at the BBC
Neuroscience Information Framework (NIF) (https://web.archive.org/web/20070125124755/h
ttp://www.neuinfo.org/)
Neurobiology (https://curlie.org/Science/Biology/Neurobiology/) at Curlie
American Society for Neurochemistry (http://www.asneurochem.org/)
British Neuroscience Association (BNA) (http://www.bna.org.uk/)
Federation of European Neuroscience Societies (http://www.fens.org/)
Neuroscience Online (electronic neuroscience textbook) (http://nba.uth.tmc.edu/neuroscienc
e/)
HHMI Neuroscience lecture series - Making Your Mind: Molecules, Motion, and Memory (htt
p://www.hhmi.org/biointeractive/neuroscience/lectures.html) Archived (https://web.archive.or
g/web/20130624074247/http://www.hhmi.org/biointeractive/neuroscience/lectures.html)
2013-06-24 at the Wayback Machine
Société des Neurosciences (http://www.neurosciences.asso.fr/)
Neuroscience For Kids (http://faculty.washington.edu/chudler/neurok.html)
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Neuroscience

Neuroscience is the scientific study of the nervous system

The scope of neuroscience has broadened over time to

Early views on the function of the brain regarded it to be a

In the process of treating epilepsy, Wilder Penfield

The understanding of neurons and of nervous

Molecular and cellular neuroscience

Questions addressed in cellular neuroscience include the mechanisms

Neural circuits and systems

Cognitive and behavioral neuroscience

Behavior of nanomaterials are dominated by long ranged non-bonding interactions.[66] Electrochemical

Neuroscience and medicine

Integrative neuroscience describes the effort to combine models and

Affective neuroscience is the study of the neural mechanisms involved in emotion,

Evolutionary neuroscience Evolutionary neuroscience studies the evolution of nervous systems.[82]

Nanoneuroscience An interdisciplinary field that integrates nanotechnology and neuroscience.[84]

Neural engineering uses engineering techniques to interact with, understand, repair,

Neuroanatomy Neuroanatomy is the study of the anatomy of nervous systems.[86]

Careers in Neuroscience [101]

Pharmaceutical Sales Residential Counselor

Laboratory Technician Regulatory Affairs Specialist

Science Writer Clinical Research Assistant

Science Advocacy Special Education Assistant

Health Educator Orthotic and Prosthetic Technician*

EEG Technologist* Lab Animal Care Technician

Forensic Science Technician Law Enforcement

Pharmacy Technician* Natural Sciences Manager

Nurse Practitioner Neuroimaging Technician

Physician's Assistant Teacher

Genetic Counselor Epidemiology

Orthotist/Prosthetist Speech-Language Pathologist

Neural Engineer Public Health

Medicine (MD, DO) Food Scientist

Research Scientist Pharmacist

Clinical Psychologist Professor

Public education and outreach

Engineering applications of neuroscience

Neuromorphic computer chips

One example of such a computer is the SpiNNaker supercomputer.[123]

Nobel prizes related to neuroscience

"for his work on

"for his work on

John Carew 1903–

1967 Physiology "for their [140]

Karl von 1886–

1981 Physiology "for his

Rita Levi- 1909–

"for the first

2003 Chemistry Roderick 1956– United States "for discoveries [147]

John United States

2017 Physiology "for their [151]

David Julius 1955– United States "for their

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