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Neuroscience is the scientific study of the
nervous system In biology, the nervous system is the complex system, highly complex part of an animal that coordinates its behavior, actions and sense, sensory information by transmitting action potential, signals to and from different parts of its body. Th ...
(the
brain The brain is an organ (biology), organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It consists of nervous tissue and is typically located in the head (cephalization), usually near organs for ...
,
spinal cord The spinal cord is a long, thin, tubular structure made up of nervous tissue that extends from the medulla oblongata in the lower brainstem to the lumbar region of the vertebral column (backbone) of vertebrate animals. The center of the spinal c ...
, and
peripheral nervous system The peripheral nervous system (PNS) is one of two components that make up the nervous system of Bilateria, bilateral animals, with the other part being the central nervous system (CNS). The PNS consists of nerves and ganglia, which lie outside t ...
), its functions, and its disorders. It is a
multidisciplinary An academic discipline or academic field is a subdivision of knowledge that is taught and researched at the college or university level. Disciplines are defined (in part) and recognized by the academic journals in which research is published, ...
science that combines
physiology Physiology (; ) is the science, scientific study of function (biology), functions and mechanism (biology), mechanisms in a life, living system. As a branches of science, subdiscipline of biology, physiology focuses on how organisms, organ syst ...
,
anatomy Anatomy () is the branch of morphology concerned with the study of the internal structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old scien ...
,
molecular biology Molecular biology is a branch of biology that seeks to understand the molecule, molecular basis of biological activity in and between Cell (biology), cells, including biomolecule, biomolecular synthesis, modification, mechanisms, and interactio ...
,
developmental biology Developmental biology is the study of the process by which animals and plants grow and develop. Developmental biology also encompasses the biology of Regeneration (biology), regeneration, asexual reproduction, metamorphosis, and the growth and di ...
, cytology,
psychology Psychology is the scientific study of mind and behavior. Its subject matter includes the behavior of humans and nonhumans, both consciousness, conscious and Unconscious mind, unconscious phenomena, and mental processes such as thoughts, feel ...
,
physics Physics is the scientific study of matter, its Elementary particle, fundamental constituents, its motion and behavior through space and time, and the related entities of energy and force. "Physical science is that department of knowledge whi ...
,
computer science Computer science is the study of computation, information, and automation. Computer science spans Theoretical computer science, theoretical disciplines (such as algorithms, theory of computation, and information theory) to Applied science, ...
,
chemistry Chemistry is the scientific study of the properties and behavior of matter. It is a physical science within the natural sciences that studies the chemical elements that make up matter and chemical compound, compounds made of atoms, molecules a ...
,
medicine Medicine is the science and Praxis (process), practice of caring for patients, managing the Medical diagnosis, diagnosis, prognosis, Preventive medicine, prevention, therapy, treatment, Palliative care, palliation of their injury or disease, ...
,
statistics Statistics (from German language, German: ', "description of a State (polity), state, a country") is the discipline that concerns the collection, organization, analysis, interpretation, and presentation of data. In applying statistics to a s ...
, and
mathematical modeling A mathematical model is an abstract and concrete, abstract description of a concrete system using mathematics, mathematical concepts and language of mathematics, language. The process of developing a mathematical model is termed ''mathematical m ...
to understand the fundamental and emergent properties of
neuron A neuron (American English), neurone (British English), or nerve cell, is an membrane potential#Cell excitability, excitable cell (biology), cell that fires electric signals called action potentials across a neural network (biology), neural net ...
s, glia and neural circuits. The understanding of the biological basis of
learning Learning is the process of acquiring new understanding, knowledge, behaviors, skills, value (personal and cultural), values, Attitude (psychology), attitudes, and preferences. The ability to learn is possessed by humans, non-human animals, and ...
,
memory Memory is the faculty of the mind by which data or information is encoded, stored, and retrieved when needed. It is the retention of information over time for the purpose of influencing future action. If past events could not be remembe ...
,
behavior Behavior (American English) or behaviour (British English) is the range of actions of Individual, individuals, organisms, systems or Artificial intelligence, artificial entities in some environment. These systems can include other systems or or ...
,
perception Perception () is the organization, identification, and interpretation of sensory information in order to represent and understand the presented information or environment. All perception involves signals that go through the nervous syste ...
, and
consciousness Consciousness, at its simplest, is awareness of a state or object, either internal to oneself or in one's external environment. However, its nature has led to millennia of analyses, explanations, and debate among philosophers, scientists, an ...
has been described by Eric Kandel as the "epic challenge" of the biological sciences. 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
neuroscientist A neuroscientist (or neurobiologist) is a scientist specializing in neuroscience that deals with the anatomy and function of neurons, Biological neural network, neural circuits, and glia, and their Behavior, behavioral, biological, and psycholo ...
s have expanded enormously, from molecular 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 Ancient Egypt () was a cradle of civilization concentrated along the lower reaches of the Nile River in Northeast Africa. It emerged from prehistoric Egypt around 3150BC (according to conventional Egyptian chronology), when Upper and Lower E ...
.
Trepanation Trepanning, also known as trepanation, trephination, trephining or making a burr hole (the verb ''trepan'' derives from Old French from Medieval Latin from Ancient Greek, Greek , literally "borer, auger"), is a surgical intervention in which a ...
, 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 The Neolithic or New Stone Age (from Ancient Greek, Greek 'new' and 'stone') is an archaeological period, the final division of the Stone Age in Mesopotamia, Asia, Europe and Africa (c. 10,000 BCE to c. 2,000 BCE). It saw the Neolithic Revo ...
period. Manuscripts dating to 1700 BC indicate that the
Egyptians Egyptians (, ; , ; ) are an ethnic group native to the Nile, Nile Valley in Egypt. Egyptian identity is closely tied to Geography of Egypt, geography. The population is concentrated in the Nile Valley, a small strip of cultivable land stretchi ...
had some knowledge about symptoms of brain damage. Early views on the function of the brain regarded it to be a "cranial stuffing" of sorts. In
Egypt Egypt ( , ), officially the Arab Republic of Egypt, is a country spanning the Northeast Africa, northeast corner of Africa and Western Asia, southwest corner of Asia via the Sinai Peninsula. It is bordered by the Mediterranean Sea to northe ...
, from the late Middle Kingdom onwards, the brain was regularly removed in preparation for mummification. It was believed at the time that the
heart The heart is a muscular Organ (biology), organ found in humans and other animals. This organ pumps blood through the blood vessels. The heart and blood vessels together make the circulatory system. The pumped blood carries oxygen and nutrie ...
was the seat of intelligence. According to
Herodotus Herodotus (; BC) was a Greek historian and geographer from the Greek city of Halicarnassus (now Bodrum, Turkey), under Persian control in the 5th century BC, and a later citizen of Thurii in modern Calabria, Italy. He wrote the '' Histori ...
, 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." The view that the heart was the source of consciousness was not challenged until the time of the Greek physician
Hippocrates Hippocrates of Kos (; ; ), also known as Hippocrates II, was a Greek physician and philosopher of the Classical Greece, classical period who is considered one of the most outstanding figures in the history of medicine. He is traditionally referr ...
. 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.
Plato Plato ( ; Greek language, Greek: , ; born  BC, died 348/347 BC) was an ancient Greek philosopher of the Classical Greece, Classical period who is considered a foundational thinker in Western philosophy and an innovator of the writte ...
also speculated that the brain was the seat of the rational part of the soul.
Aristotle Aristotle (; 384–322 BC) was an Ancient Greek philosophy, Ancient Greek philosopher and polymath. His writings cover a broad range of subjects spanning the natural sciences, philosophy, linguistics, economics, politics, psychology, a ...
, however, believed the heart was the center of intelligence and that the brain regulated the amount of heat from the heart. This view was generally accepted until the Roman physician
Galen Aelius Galenus or Claudius Galenus (; September 129 – AD), often Anglicization, anglicized as Galen () or Galen of Pergamon, was a Ancient Rome, Roman and Greeks, Greek physician, surgeon, and Philosophy, philosopher. Considered to be one o ...
, a follower of Hippocrates and physician to
Roman gladiators A gladiator ( , ) was an armed combatant who entertained audiences in the Roman Republic and Roman Empire in violent confrontations with other gladiators, wild animals, and condemned criminals. Some gladiators were volunteers who risked their ...
, observed that his patients lost their mental faculties when they had sustained damage to their brains. Abulcasis, Averroes,
Avicenna Ibn Sina ( – 22 June 1037), commonly known in the West as Avicenna ( ), was a preeminent philosopher and physician of the Muslim world, flourishing during the Islamic Golden Age, serving in the courts of various Iranian peoples, Iranian ...
, Avenzoar, and
Maimonides Moses ben Maimon (1138–1204), commonly known as Maimonides (, ) and also referred to by the Hebrew acronym Rambam (), was a Sephardic rabbi and Jewish philosophy, philosopher who became one of the most prolific and influential Torah schola ...
, active in the Medieval Muslim world, described a number of medical problems related to the brain. In Renaissance Europe,
Vesalius Andries van Wezel (31 December 1514 – 15 October 1564), Latinization of names, latinized as Andreas Vesalius (), was an anatomist and physician who wrote ''De humani corporis fabrica, De Humani Corporis Fabrica Libri Septem'' (''On the fabric ...
(1514–1564),
René Descartes René Descartes ( , ; ; 31 March 1596 – 11 February 1650) was a French philosopher, scientist, and mathematician, widely considered a seminal figure in the emergence of modern philosophy and Modern science, science. Mathematics was paramou ...
(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, whose speed Hermann von Helmholtz proceeded to measure, and in 1875 Richard Caton found electrical phenomena in the cerebral hemispheres of rabbits and monkeys. Adolf Beck published in 1890 similar observations of spontaneous electrical activity of the brain of rabbits and dogs. Studies of the brain became more sophisticated after the invention of the
microscope A microscope () is a laboratory equipment, laboratory instrument used to examine objects that are too small to be seen by the naked eye. Microscopy is the science of investigating small objects and structures using a microscope. Microscopic ...
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
neuron A neuron (American English), neurone (British English), or nerve cell, is an membrane potential#Cell excitability, excitable cell (biology), cell that fires electric signals called action potentials across a neural network (biology), neural net ...
s. 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. 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 The cerebral cortex, also known as the cerebral mantle, is the outer layer of neural tissue of the cerebrum of the brain in humans and other mammals. It is the largest site of Neuron, neural integration in the central nervous system, and plays ...
. 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 The motor cortex is the region of the cerebral cortex involved in the planning, motor control, control, and execution of voluntary movements. The motor cortex is an area of the frontal lobe located in the posterior precentral gyrus immediately ...
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 Neuroimaging is the use of quantitative (computational) techniques to study the neuroanatomy, structure and function of the central nervous system, developed as an objective way of scientifically studying the healthy human brain in a non-invasive ...
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. 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. 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 The Massachusetts Institute of Technology (MIT) is a Private university, private research university in Cambridge, Massachusetts, United States. Established in 1861, MIT has played a significant role in the development of many areas of moder ...
, 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. This was followed by the Department of Neurobiology at Harvard Medical School, which was founded in 1966 by Stephen Kuffler. In the process of treating
epilepsy Epilepsy is a group of Non-communicable disease, non-communicable Neurological disorder, neurological disorders characterized by a tendency for recurrent, unprovoked Seizure, seizures. A seizure is a sudden burst of abnormal electrical activit ...
, Wilder Penfield produced maps of the location of various functions (motor, sensory, memory, vision) in the brain. He summarized his findings in a 1950 book called ''The Cerebral Cortex of Man''. Wilder Penfield and his co-investigators Edwin Boldrey and Theodore Rasmussen are considered to be the originators of the cortical homunculus. 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 A mathematical model is an abstract and concrete, abstract description of a concrete system using mathematics, mathematical concepts and language of mathematics, language. The process of developing a mathematical model is termed ''mathematical m ...
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, the International Society for Neurochemistry in 1963, the European Brain and Behaviour Society in 1968, and the Society for Neuroscience in 1969. Recently, the application of neuroscience research results has also given rise to applied disciplines as
neuroeconomics Neuroeconomics is an Interdisciplinarity, interdisciplinary field that seeks to explain human decision-making, the ability to process multiple alternatives and to follow through on a plan of action. It studies how economic behavior can shape our u ...
, neuroeducation, neuroethics, and neurolaw. 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.


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 Molecular biology is a branch of biology that seeks to understand the molecule, molecular basis of biological activity in and between Cell (biology), cells, including biomolecule, biomolecular synthesis, modification, mechanisms, and interactio ...
,
electrophysiology Electrophysiology (from ee the Electron#Etymology, etymology of "electron" ; and ) is the branch of physiology that studies the electrical properties of biological cell (biology), cells and tissues. It involves measurements of voltage change ...
, and computational neuroscience. This has allowed neuroscientists to study the
nervous system In biology, the nervous system is the complex system, highly complex part of an animal that coordinates its behavior, actions and sense, sensory information by transmitting action potential, signals to and from different parts of its body. Th ...
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 A neuron (American English), neurone (British English), or nerve cell, is an membrane potential#Cell excitability, excitable cell (biology), cell that fires electric signals called action potentials across a neural network (biology), neural net ...
. 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 An axon (from Greek ἄξων ''áxōn'', axis) or nerve fiber (or nerve fibre: see American and British English spelling differences#-re, -er, spelling differences) is a long, slender cellular extensions, projection of a nerve cell, or neuron, ...
, 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 parts: the
central nervous system The central nervous system (CNS) is the part of the nervous system consisting primarily of the brain, spinal cord and retina. The CNS is so named because the brain integrates the received information and coordinates and influences the activity o ...
(defined as the
brain The brain is an organ (biology), organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It consists of nervous tissue and is typically located in the head (cephalization), usually near organs for ...
and
spinal cord The spinal cord is a long, thin, tubular structure made up of nervous tissue that extends from the medulla oblongata in the lower brainstem to the lumbar region of the vertebral column (backbone) of vertebrate animals. The center of the spinal c ...
), and the
peripheral nervous system The peripheral nervous system (PNS) is one of two components that make up the nervous system of Bilateria, bilateral animals, with the other part being the central nervous system (CNS). The PNS consists of nerves and ganglia, which lie outside t ...
. 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 The human brain is the central organ (anatomy), organ of the nervous system, and with the spinal cord, comprises the central nervous system. It consists of the cerebrum, the brainstem and the cerebellum. The brain controls most of the activi ...
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. Due to the high degree of plasticity of the human brain, the structure of its synapses and their resulting functions change throughout life. 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 Computer science is the study of computation, information, and automation. Computer science spans Theoretical computer science, theoretical disciplines (such as algorithms, theory of computation, and information theory) to Applied science, ...
,
electronics Electronics is a scientific and engineering discipline that studies and applies the principles of physics to design, create, and operate devices that manipulate electrons and other Electric charge, electrically charged particles. It is a subfield ...
,
functional neuroimaging Functional neuroimaging is the use of neuroimaging technology to measure an aspect of brain function, often with a view to understanding the relationship between activity in certain brain areas and specific mental functions. It is primarily used a ...
, and
genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinians, Augustinian ...
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. 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.


Molecular and cellular neuroscience

Basic questions addressed in molecular neuroscience include the mechanisms by which neurons express and respond to molecular signals and how
axon An axon (from Greek ἄξων ''áxōn'', axis) or nerve fiber (or nerve fibre: see American and British English spelling differences#-re, -er, spelling differences) is a long, slender cellular extensions, projection of a nerve cell, or neuron, ...
s form complex connectivity patterns. At this level, tools from
molecular biology Molecular biology is a branch of biology that seeks to understand the molecule, molecular basis of biological activity in and between Cell (biology), cells, including biomolecule, biomolecular synthesis, modification, mechanisms, and interactio ...
and
genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinians, Augustinian ...
are used to understand how neurons develop and how genetic changes affect biological functions. The morphology, molecular identity, and physiological characteristics of neurons and how they relate to different types of behavior are also of considerable interest. Questions addressed in
cellular neuroscience Cellular neuroscience is a branch of neuroscience concerned with the study of neurons at a cellular level. This includes Morphology (biology), morphology and physiology, physiological properties of single neurons. Several techniques such as intrace ...
include the mechanisms of how neurons process signals physiologically and electrochemically. These questions include how signals are processed by neurites and somas and how
neurotransmitter A neurotransmitter is a signaling molecule secreted by a neuron to affect another cell across a Chemical synapse, synapse. The cell receiving the signal, or target cell, may be another neuron, but could also be a gland or muscle cell. Neurotra ...
s 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
axon An axon (from Greek ἄξων ''áxōn'', axis) or nerve fiber (or nerve fibre: see American and British English spelling differences#-re, -er, spelling differences) is a long, slender cellular extensions, projection of a nerve cell, or neuron, ...
s (specialized to conduct nerve impulses called
action potential An action potential (also known as a nerve impulse or "spike" when in a neuron) is a series of quick changes in voltage across a cell membrane. An action potential occurs when the membrane potential of a specific Cell (biology), cell rapidly ri ...
s). 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 The development of the nervous system, or neural development (neurodevelopment), refers to the processes that generate, shape, and reshape the nervous system of animals, from the earliest stages of embryonic development to adulthood. The field ...
. 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, differentiation of neurons and glia ( neurogenesis and gliogenesis), and neuronal migration. 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 (Computational Neurogenetic Modeling (CNGM) can also be used to model neural systems).


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 network Large-scale brain networks (also known as intrinsic brain networks) are collections of widespread brain regions showing Resting state fMRI#Functional, functional connectivity by statistical analysis of the Functional magnetic resonance imaging, fMR ...
s, 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. Questions in systems neuroscience include how neural circuits are formed and used anatomically and physiologically to produce functions such as reflexes,
multisensory integration Multisensory integration, also known as multimodal integration, is the study of how information from the different sensory modality, sensory modalities (such as sight, sound, touch, smell, self-motion, and taste) may be integrated by the nervous sy ...
,
motor coordination In physiology, motor coordination is the orchestrated movement of multiple body parts as required to accomplish intended actions, like walking. This coordination is achieved by adjusting kinematic and kinetic parameters associated with each bo ...
,
circadian rhythm A circadian rhythm (), or circadian cycle, is a natural oscillation that repeats roughly every 24 hours. Circadian rhythms can refer to any process that originates within an organism (i.e., Endogeny (biology), endogenous) and responds to the env ...
s, emotional responses,
learning Learning is the process of acquiring new understanding, knowledge, behaviors, skills, value (personal and cultural), values, Attitude (psychology), attitudes, and preferences. The ability to learn is possessed by humans, non-human animals, and ...
, and
memory Memory is the faculty of the mind by which data or information is encoded, stored, and retrieved when needed. It is the retention of information over time for the purpose of influencing future action. If past events could not be remembe ...
. 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. 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. The related fields of neuroethology and
neuropsychology Neuropsychology is a branch of psychology concerned with how a person's cognition and behavior are related to the brain and the rest of the nervous system. Professionals in this branch of psychology focus on how injuries or illnesses of the brai ...
address the question of how neural substrates underlie specific
animal Animals are multicellular, eukaryotic organisms in the Biology, biological Kingdom (biology), kingdom Animalia (). With few exceptions, animals heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, ...
and
human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
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 Neuroimaging is the use of quantitative (computational) techniques to study the neuroanatomy, structure and function of the central nervous system, developed as an objective way of scientifically studying the healthy human brain in a non-invasive ...
(e.g., fMRI, PET, SPECT), EEG, MEG,
electrophysiology Electrophysiology (from ee the Electron#Etymology, etymology of "electron" ; and ) is the branch of physiology that studies the electrical properties of biological cell (biology), cells and tissues. It involves measurements of voltage change ...
,
optogenetics Optogenetics is a biological technique to control the activity of neurons or other cell types with light. This is achieved by Gene expression, expression of Channelrhodopsin, light-sensitive ion channels, Halorhodopsin, pumps or Photoactivated ade ...
and human genetic analysis combined with sophisticated experimental techniques from
cognitive psychology Cognitive psychology is the scientific study of human mental processes such as attention, language use, memory, perception, problem solving, creativity, and reasoning. Cognitive psychology originated in the 1960s in a break from behaviorism, whi ...
allows
neuroscientist A neuroscientist (or neurobiologist) is a scientist specializing in neuroscience that deals with the anatomy and function of neurons, Biological neural network, neural circuits, and glia, and their Behavior, behavioral, biological, and psycholo ...
s and
psychologist A psychologist is a professional who practices psychology and studies mental states, perceptual, cognitive, emotional, and social processes and behavior. Their work often involves the experimentation, observation, and explanation, interpretatio ...
s to address abstract questions such as how cognition and emotion are mapped to specific neural substrates. Although many studies hold a reductionist stance looking for the neurobiological basis of cognitive phenomena, recent research shows that there is an interplay between neuroscientific findings and conceptual research, soliciting and integrating both perspectives. For example, neuroscience research on empathy solicited an interdisciplinary debate involving philosophy, psychology and psychopathology. Moreover, the neuroscientific identification of multiple memory systems related to different brain areas has challenged the idea of
memory Memory is the faculty of the mind by which data or information is encoded, stored, and retrieved when needed. It is the retention of information over time for the purpose of influencing future action. If past events could not be remembe ...
as a literal reproduction of the past, supporting a view of memory as a generative, constructive and dynamic process. Neuroscience is also allied with the
social Social organisms, including human(s), live collectively in interacting populations. This interaction is considered social whether they are aware of it or not, and whether the exchange is voluntary or not. Etymology The word "social" derives fro ...
and
behavioral sciences Behavioural science is the branch of science concerned with human behaviour.Hallsworth, M. (2023). A manifesto for applying behavioural science. ''Nature Human Behaviour'', ''7''(3), 310-322. While the term can technically be applied to the st ...
, as well as with nascent interdisciplinary fields. Examples of such alliances include
neuroeconomics Neuroeconomics is an Interdisciplinarity, interdisciplinary field that seeks to explain human decision-making, the ability to process multiple alternatives and to follow through on a plan of action. It studies how economic behavior can shape our u ...
,
decision theory Decision theory or the theory of rational choice is a branch of probability theory, probability, economics, and analytic philosophy that uses expected utility and probabilities, probability to model how individuals would behave Rationality, ratio ...
, 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.


Computational neuroscience

Questions in computational neuroscience can span a wide range of levels of traditional analysis, such as development,
structure A structure is an arrangement and organization of interrelated elements in a material object or system, or the object or system so organized. Material structures include man-made objects such as buildings and machines and natural objects such as ...
, 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 A neural network is a group of interconnected units called neurons that send signals to one another. Neurons can be either Cell (biology), biological cells or signal pathways. While individual neurons are simple, many of them together in a netwo ...
. Computational neuroscience is often referred to as theoretical neuroscience.


Neuroscience and medicine


Clinical neuroscience

Neurology, psychiatry, neurosurgery, psychosurgery, anesthesiology and pain medicine, neuropathology, neuroradiology,
ophthalmology Ophthalmology (, ) is the branch of medicine that deals with the diagnosis, treatment, and surgery of eye diseases and disorders. An ophthalmologist is a physician who undergoes subspecialty training in medical and surgical eye care. Following a ...
, 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.
Neurology Neurology (from , "string, nerve" and the suffix wikt:-logia, -logia, "study of") is the branch of specialty (medicine) , medicine dealing with the diagnosis and treatment of all categories of conditions and disease involving the nervous syst ...
works with diseases of the central and peripheral nervous systems, such as
amyotrophic lateral sclerosis Amyotrophic lateral sclerosis (ALS), also known as motor neuron disease (MND) or—in the United States—Lou Gehrig's disease (LGD), is a rare, Terminal illness, terminal neurodegenerative disease, neurodegenerative disorder that results i ...
(ALS) and
stroke Stroke is a medical condition in which poor cerebral circulation, blood flow to a part of the brain causes cell death. There are two main types of stroke: brain ischemia, ischemic, due to lack of blood flow, and intracranial hemorrhage, hemor ...
, and their medical treatment.
Psychiatry Psychiatry is the medical specialty devoted to the diagnosis, treatment, and prevention of deleterious mental disorder, mental conditions. These include matters related to cognition, perceptions, Mood (psychology), mood, emotion, and behavior. ...
focuses on affective, behavioral, cognitive, and perceptual disorders.
Anesthesiology Anesthesiology, anaesthesiology or anaesthesia is the medical specialty concerned with the total perioperative medicine, perioperative care of patients before, during and after surgery. It encompasses anesthesia, intensive care medicine, critica ...
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 Neurosurgery or neurological surgery, known in common parlance as brain surgery, is the specialty (medicine), medical specialty that focuses on the surgical treatment or rehabilitation of disorders which affect any portion of the nervous system ...
and psychosurgery work primarily with surgical treatment of diseases of the central and peripheral nervous systems. Neuroscience underlies the development of various neurotherapy methods to treat diseases of the nervous system.IEEE Brain (2019). "Neurotherapy: Treating Disorders by Retraining the Brain". ''The Future Neural Therapeutics White Paper''. Retrieved 23.01.2025 from: https://brain.ieee.org/topics/neurotherapy-treating-disorders-by-retraining-the-brain/#:~:text=Neurotherapy%20trains%20a%20patient's%20brain,wave%20activity%20through%20positive%20reinforcement International Neuromodulation Society, Retrieved 23 January 2025 from: https://www.neuromodulation.com/ Val Danilov I (2023). "The Origin of Natural Neurostimulation: A Narrative Review of Noninvasive Brain Stimulation Techniques." ''OBM Neurobiology'' 2024; 8(4): 260; https://doi:10.21926/obm.neurobiol.2404260.


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. 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. Another important area of translational research is
brain–computer interface A brain–computer interface (BCI), sometimes called a brain–machine interface (BMI), is a direct communication link between the brain's electrical activity and an external device, most commonly a computer or robotic limb. BCIs are often dire ...
s (BCIs), or machines that are able to communicate and influence the brain. They are currently being researched for their potential to repair neural systems and restore certain cognitive functions. However, some ethical considerations have to be dealt with before they are accepted.


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.


Careers in neuroscience

Source:


Bachelor's Level


Master's Level


Advanced Degree


Neuroscience organizations

The largest professional neuroscience organization is the Society for Neuroscience (SFN), which is based in the
United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
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. 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 (), and the French '. 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 and Project Encephalon. In 2013, the BRAIN Initiative was announced in the US. The International Brain Initiative was created in 2017, currently integrated by more than seven national-level brain research initiatives (US,
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
, Allen Institute,
Japan Japan is an island country in East Asia. Located in the Pacific Ocean off the northeast coast of the Asia, Asian mainland, it is bordered on the west by the Sea of Japan and extends from the Sea of Okhotsk in the north to the East China Sea ...
,
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
, Australia, Canada, Korea, and Israel) 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. In the United States, large organizations such as the Society for Neuroscience have promoted neuroscience education by developing a primer called Brain Facts, collaborating with public school teachers to develop Neuroscience Core Concepts for K-12 teachers and students, and cosponsoring a campaign with the Dana Foundation called Brain Awareness Week to increase public awareness about the progress and benefits of brain research. In Canada, the Canadian Institutes of Health Research's (CIHR) Canadian National Brain Bee is held annually at McMaster University. Neuroscience educators formed a Faculty for Undergraduate Neuroscience (FUN) in 1992 to share best practices and provide travel awards for undergraduates presenting at Society for Neuroscience meetings. 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. Federal agencies in the United States, such as the National Institute of Health (NIH) and
National Science Foundation The U.S. National Science Foundation (NSF) is an Independent agencies of the United States government#Examples of independent agencies, independent agency of the Federal government of the United States, United States federal government that su ...
(NSF), 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
complex systems A complex system is a system composed of many components that may interact with one another. Examples of complex systems are Earth's global climate, organisms, the human brain, infrastructure such as power grid, transportation or communication s ...
, and that the computational components are interrelated with no central processor. One example of such a computer is the SpiNNaker supercomputer. 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. Recent advances in neuromorphic microchip technology have led a group of scientists to create an artificial neuron that can replace real neurons in diseases.


Nobel prizes related to neuroscience


See also

* List of neuroscience databases * List of neuroscience journals * List of neuroscientists * Neurosemiotics * Neuroscience of sex differences * Outline of brain mapping * Outline of the human brain * List of regions in the human brain * Gut–brain axis * Connectomics


References


Further reading

* * * Squire, L. ''et al.'' (2012). ''Fundamental Neuroscience, 4th edition''. Academic Press; * Byrne and Roberts (2004). ''From Molecules to Networks''. Academic Press; * Sanes, Reh, Harris (2005). ''Development of the Nervous System, 2nd edition''. Academic Press; * Siegel ''et al.'' (2005). ''Basic Neurochemistry, 7th edition''. Academic Press; * Rieke, F. ''et al.'' (1999). ''Spikes: Exploring the Neural Code''. The MIT Press; Reprint edition
section.47 Neuroscience
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
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. * * Damasio, A. R. (1994). ''Descartes' Error: Emotion, Reason, and the Human Brain. '' New York, Avon (publishers), Avon Books. (Hardcover) (Paperback) * Gardner, H. (1976). ''The Shattered Mind: The Person After Brain Damage. '' New York, Random House, Vintage Books, 1976 * Goldstein, K. (2000). ''The Organism. '' New York, Zone Books. (Hardcover) (Paperback) * * Subhash Kak, The Architecture of Knowledge: Quantum Mechanics, Neuroscience, Computers and Consciousness, Motilal Banarsidass, 2004, * Llinas R. (2001). ''I of the vortex: from neurons to self'' MIT Press. (Hardcover) (Paperback) * Luria, A. R. (1997). ''The Man with a Shattered World: The History of a Brain Wound. '' Cambridge, Massachusetts, Harvard University Press. (Hardcover) (Paperback) * Luria, A. R. (1998). ''The Mind of a Mnemonist: A Little Book About A Vast Memory. '' New York, Basic Books, Inc. * Medina, J. (2008). ''Brain Rules: 12 Principles for Surviving and Thriving at Work, Home, and School. ''Seattle, Pear Press. (Hardcover with DVD) * Pinker, S. (1999). ''How the Mind Works. '' W. W. Norton & Company. * Pinker, S. (2002). ''The Blank Slate: The Modern Denial of Human Nature. '' Viking Adult. * * 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. (Paperback) * Rose, S. (2006). ''21st Century Brain: Explaining, Mending & Manipulating the Mind'' (Paperback) * Sacks, O. ''The Man Who Mistook His Wife for a Hat''. Summit Books (Hardcover) (Paperback) * Sacks, O. (1990). ''Awakenings. '' New York, Vintage Books. (See also Oliver Sacks) (Hardcover) (Paperback)
Encyclopedia:Neuroscience
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''
. Princeton University Press. *


External links

*
Neuroscience Information Framework (NIF)

American Society for Neurochemistry

British Neuroscience Association (BNA)

Federation of European Neuroscience Societies

Neuroscience Online (electronic neuroscience textbook)


* [http://www.neurosciences.asso.fr/ ''Société des Neurosciences'']
Neuroscience For Kids
{{Authority control Neuroscience, Neurology Nervous system Neurophysiology