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Neuroscience (or neurobiology) is the scientific study of the nervous system.[1] It is a multidisciplinary branch of biology[2] that combines physiology, anatomy, molecular biology, developmental biology, cytology, mathematical modeling and psychology to understand the fundamental and emergent properties of neurons and neural circuits.[3][4][5][6][7] The understanding of the biological basis of learning, memory, behavior, perception, and consciousness has been described by Eric Kandel as the "ultimate challenge" of the biological sciences.[8]

The scope of neuroscience has broadened over time to include different approaches used to study the nervous system at different scales and the techniques used by neuroscientists have expanded enormously, from molecular and cellular studies of individual neurons to imaging of sensory, motor and cognitive tasks in the brain.



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.[9]



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.

The Golgi stain first allowed for the visualization of individual neurons.

Luigi Galvani's pioneering work in the late 1700s set the stage for studying the electrical excitability of muscles and neurons. In the first half of the 19th century, Jean Pierre Flourens pioneered the experimental method of carrying out localized lesions of the brain in living animals describing their effects on motricity, sensibility and behavior. In 1843 Emil du Bois-Reymond demonstrated the electrical nature of the nerve signal,[13] whose speed Hermann von Helmholtz proceeded to measure,[14] and in 1875 Richard Caton found electrical phenomena in the cerebral hemispheres of rabbits and monkeys.[15] Adolf Beck published in 1890 similar observations of spontaneous electrical activity of the brain of rabbits and dogs.[16] Studies of the brain became more sophisticated after the invention of the 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.[17] 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, work with brain-damaged patients by Paul Broca suggested that certain regions of the brain were responsible for certain functions. At the time, Broca's 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.[18][19] 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 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.[20]

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.[21] 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.[22] This was followed by the Department of Neurobiology at Harvard Medical School, which was founded in 1966 by Stephen Kuffler.[23]

The understanding of neurons and of nervous system function became increasingly precise and molecular during the 20th century. For example, 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, 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 neuroscientist during the 20th century. For example, the International Brain Research Organization was founded in 1961,[24] the International Society for Neurochemistry in 1963,[25] the European Brain and Behaviour Society in 1968,[26] and the Society for Neuroscience in 1969.[27] Recently, the application of neuroscience research results has also given rise to applied disciplines as neuroeconomics,[28] neuroeducation,[29] neuroethics,[30] and neurolaw.[31]

Over time, brain research has gone through philosophical, experimental, and theoretical phases, with work on brain simulation predicted to be important in the future.[32]

Modern neuroscience

Human nervous system



The vertebrate nervous system can be split into two 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.[33]

Due to the high degree of plasticity of the human brain, the structure of its synapses and their resulting functions change throughout life.[34]

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 foci 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.

Molecular and cellular neuroscience

Photograph of a stained neuron in a chicken embryo


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 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.

Another major area of cellular neuroscience is the investigation of the development of the nervous system. Questions include the patterning and regionalization of the nervous system, neural stem cells, differentiation of neurons and glia (neurogenesis and gliogenesis), neuronal migration, axonal and dendritic development, trophic interactions, and synapse formation.

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.

Neural circuits and systems

Proposed organization of motor-semantic neural circuits for action language comprehension. Adapted from Shebani et al. (2013)

Questions in systems neuroscience include how neural circuits are formed and used anatomically and physiologically to produce functions such as reflexes, multisensory integration, motor coordination, circadian rhythms, emotional responses, learning, and memory. In other words, they address how these neural circuits function in large scale brain networks, and the mechanisms through which behaviors are generated. For example, systems level analysis addresses questions concerning specific sensory and motor modalities: how does vision work? How do songbirds learn new songs and bats localize with ultrasound? How does the somatosensory system process tactile information? The related fields of neuroethology and neuropsychology address the question of how neural substrates underlie specific animal and human behaviors. Neuroendocrinology and psychoneuroimmunology examine interactions between the nervous system and the endocrine and immune systems, respectively. Despite many advancements, the way that networks of neurons perform complex cognitive processes and behaviors is still poorly understood.

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, the neuroscience research on empathy solicited an interesting interdisciplinary debate involving philosophy, psychology and psychopathology.[35] 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.[36]

Neuroscience is also allied with the social and behavioral sciences as well as nascent interdisciplinary fields such as 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.[37]

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.

Translational research and medicine

Parasagittal MRI of the head of a patient with benign familial macrocephaly

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.


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.[38]

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.[39]

Major branches


List of the major branches of neuroscience
Branch Description
Behavioral neuroscience Behavioral neuroscience (also known as biological psychology, physiological psychology, biopsychology, or psychobiology) is the application of the principles of biology to the study of genetic, physiological, and developmental mechanisms of behavior in humans and non-human animals.
Clinical neuroscience The scientific study of the biological mechanisms that underlie the disorders and diseases of the nervous system.
Cognitive neuroscience Cognitive neuroscience is the study of the biological mechanisms underlying cognition.
Computational neuroscience Computational neuroscience is the theoretical study of the nervous system.
Evolutionary neuroscience Evolutionary neuroscience studies the evolution of nervous systems.
Molecular neuroscience Molecular neuroscience studies the nervous system with molecular biology, molecular genetics, protein chemistry, and related methodologies.
Neural engineering Neural engineering uses engineering techniques to interact with, understand, repair, replace, or enhance neural systems.
Neuroanatomy Neuroanatomy the study of the anatomy of nervous systems.
Neurochemistry Neurochemistry is the study of how neurochemicals interact and influence the function of neurons.
Neuroethology Neuroethology is the study of the neural basis of non-human animals behavior.
Neurogastronomy Neurogastronomy is the study of flavor and how it affects sensation, cognition, and memory.[42]
Neurogenetics Neurogenetics is the study of the genetical basis of the development and function of the nervous system.
Neuroimmunology Neuroimmunology is concerned with the interactions between the nervous and the immune system.
Neurophysics Neurophysics deals with the development of physical experimental tools to gain information about the brain.

Neuroscience organizations



In 2013, the BRAIN Initiative was announced in the US. An International Brain Initiative was created in 2017,[44] currently integrated by more than seven national-level brain research initiatives (US, Europe, Allen Institute, Japan, China, Australia, Canada, Korea, Israel)[45] 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.[46] In the United States, large organizations such as the Society for Neuroscience have promoted neuroscience education by developing a primer called Brain Facts,[47] collaborating with public school teachers to develop Neuroscience Core Concepts for K-12 teachers and students,[48] and cosponsoring a campaign with the Dana Foundation called Brain Awareness Week to increase public awareness about the progress and benefits of brain research.[49] In Canada, the CIHR Canadian National Brain Bee is held annually at McMaster University.[50]

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.[51]

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.[52] Federal agencies in the United States, such as the National Institute of Health (NIH)[53] and National Science Foundation (NSF),[54] have also funded research that pertains to best practices in teaching and learning of neuroscience concepts.

Nobel prizes related to Neuroscience

Year Prize field Image Laureate Lifetime Country Rationale Ref
1904 Physiology Ivan Pavlov nobel.jpg Ivan Petrovich Pavlov 1849–1936  Russia "in recognition of his work on the physiology of digestion, through which knowledge on vital aspects of the subject has been transformed and enlarged" [55]
1906 Physiology Camillo Golgi nobel.jpg Camillo Golgi 1843–1926  Italy "in recognition of their work on the structure of the nervous system" [56]
Cajal-Restored.jpg Santiago Ramón y Cajal 1852–1934  Spain
1914 Physiology Robert Barany.jpg Robert Bárány 1876–1936  Austria-Hungary "for his work on the physiology and pathology of the vestibular apparatus" [57]
1932 Physiology Prof. Charles Scott Sherrington.jpg Charles Scott Sherrington 1857–1952  United Kingdom "for their discoveries regarding the functions of neurons" [58]
Edgar Douglas Adrian nobel.jpg Edgar Douglas Adrian 1889–1977  United Kingdom
1936 Physiology Henry Dale nobel.jpg Henry Hallett Dale 1875–1968  United Kingdom "for their discoveries relating to chemical transmission of nerve impulses" [59]
Otto Loewi nobel.jpg Otto Loewi 1873–1961  Austria
1944 Physiology Joseph Erlanger nobel.jpg Joseph Erlanger 1874–1965  United States "for their discoveries relating to the highly differentiated functions of single nerve fibres" [60]
Herbert Spencer Gasser nobel.jpg Herbert Spencer Gasser 1888–1963  United States
1949 Physiology Walter Hess.jpg Walter Rudolf Hess 1881–1973   Switzerland "for his discovery of the functional organization of the interbrain as a coordinator of the activities of the internal organs" [61]
Moniz.jpg António Caetano Egas Moniz 1874–1955  Portugal "for his discovery of the therapeutic value of leucotomy in certain psychoses" [61]
1957 Physiology Daniel Bovet nobel.jpg Daniel Bovet 1907–1992  Italy "for his discoveries relating to synthetic compounds that inhibit the action of certain body substances, and especially their action on the vascular system and the skeletal muscles" [62]
1961 Physiology Georg von Békésy nobel.jpg Georg von Békésy 1899–1972  United States "for his discoveries of the physical mechanism of stimulation within the cochlea" [63]
1963 Physiology Eccles lab.jpg John Carew Eccles 1903–1997  Australia "for their discoveries concerning the ionic mechanisms involved in excitation and inhibition in the peripheral and central portions of the nerve cell membrane" [64]
Alan Lloyd Hodgkin nobel.jpg Alan Lloyd Hodgkin 1914–1998  United Kingdom
Andrew Fielding Huxley nobel.jpg Andrew Fielding Huxley 1917–2012  United Kingdom
1967 Physiology Ragnar Granit2.jpg Ragnar Granit 1900–1991  Finland
"for their discoveries concerning the primary physiological and chemical visual processes in the eye" [65]
Haldan Keffer Hartline nobel.jpg Haldan Keffer Hartline 1903–1983  United States
George Wald nobel.jpg George Wald 1906–1997  United States
1970 Physiology Julius Axelrod 1912–2004  United States "for their discoveries concerning the humoral transmittors in the nerve terminals and the mechanism for their storage, release and inactivation" [64]
Ulf von Euler.jpg Ulf von Euler 1905–1983  Sweden
Bernard Katz 1911–2003  United Kingdom
1981 Physiology Roger Wolcott Sperry.jpg Roger W. Sperry 1913–1994  United States "for his discoveries concerning the functional specialization of the cerebral hemispheres" [65]
David H. Hubel 1926–2013  Canada "for their discoveries concerning information processing in the visual system" [65]
Torsten Wiesel-7Nov2006.jpg Torsten N. Wiesel 1924–  Sweden
1986 Physiology Stanley Cohen-Biochemist.jpg Stanley Cohen 1922–  United States "for their discoveries of growth factors" [66]
Rita Levi Montalcini.jpg Rita Levi-Montalcini 1909–2012  Italy
1997 Chemistry Skou2008crop.jpg Jens C. Skou 1918–2018  Denmark "for the first discovery of an ion-transporting enzyme, Na+, K+ -ATPase" [67]
2000 Physiology Arvid Carlsson 2011a.jpg Arvid Carlsson 1923–2018  Sweden "for their discoveries concerning signal transduction in the nervous system" [68]
Paul Greengard.jpg Paul Greengard 1925–2019  United States
Eric Richard Kandel.jpg Eric R. Kandel 1929–  United States
2003 Chemistry Roderick MacKinnon Roderick MacKinnon 1956–  United States "for discoveries concerning channels in cell membranes [...] for structural and mechanistic studies of ion channels" [69]
2014 Physiology John O'Keefe (neuroscientist) 2014b.jpg John O'Keefe 1939–  United States
 United Kingdom
"for their discoveries of cells that constitute a positioning system in the brain" [70]
May-Britt Moser 2014.jpg May-Britt Moser 1963–  Norway
Edvard Moser.jpg Edvard I. Moser 1962–  Norway
2017 Physiology Jeffrey C. Hall EM1B8737 (38162359274).jpg Jeffrey C. Hall 1939–  United States "for their discoveries of molecular mechanisms controlling the circadian rhythm" [71]
Michael Rosbash EM1B8756 (38847326642).jpg Michael Rosbash 1944–  United States
Michael W. Young D81 4345 (38162439194).jpg Michael W. Young 1949–  United States

See also


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Further reading

External links