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Drawing by Santiago Ramon y Cajal (1899) of neurons in the pigeon cerebellum PurkinjeCell.jpg
Drawing by Santiago Ramón y Cajal (1899) of neurons in the pigeon cerebellum

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]

Science systematic enterprise that builds and organizes knowledge

Science is a systematic enterprise that builds and organizes knowledge in the form of testable explanations and predictions about the universe.

Nervous system Control the whole body function

The nervous system is a highly complex part of an animal that coordinates its actions and sensory information by transmitting signals to and from different parts of its body. The nervous system detects environmental changes that impact the body, then works in tandem with the endocrine system to respond to such events. Nervous tissue first arose in wormlike organisms about 550 to 600 million years ago. In vertebrates it consists of two main parts, the central nervous system (CNS) and the peripheral nervous system (PNS). The CNS consists of the brain and spinal cord. The PNS consists mainly of nerves, which are enclosed bundles of the long fibers or axons, that connect the CNS to every other part of the body. Nerves that transmit signals from the brain are called motor or efferent nerves, while those nerves that transmit information from the body to the CNS are called sensory or afferent. Spinal nerves serve both functions and are called mixed nerves. The PNS is divided into three separate subsystems, the somatic, autonomic, and enteric nervous systems. Somatic nerves mediate voluntary movement. The autonomic nervous system is further subdivided into the sympathetic and the parasympathetic nervous systems. The sympathetic nervous system is activated in cases of emergencies to mobilize energy, while the parasympathetic nervous system is activated when organisms are in a relaxed state. The enteric nervous system functions to control the gastrointestinal system. Both autonomic and enteric nervous systems function involuntarily. Nerves that exit from the cranium are called cranial nerves while those exiting from the spinal cord are called spinal nerves.

Biology is the natural science that studies life and living organisms, including their physical structure, chemical processes, molecular interactions, physiological mechanisms, development and evolution. Despite the complexity of the science, there are certain unifying concepts that consolidate it into a single, coherent field. Biology recognizes the cell as the basic unit of life, genes as the basic unit of heredity, and evolution as the engine that propels the creation and extinction of species. Living organisms are open systems that survive by transforming energy and decreasing their local entropy to maintain a stable and vital condition defined as homeostasis.


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.

Neuroscientist individual who studies neuroscience

A neuroscientist is a scientist who has specialised knowledge in the field of neuroscience, the branch of biology that deals with the physiology, biochemistry, anatomy and molecular biology of neurons and neural circuits and especially their association with behaviour and learning.

Molecular biology branch of biology that deals with the molecular basis of biological activity

Molecular biology is a branch of biology that concerns the molecular basis of biological activity between biomolecules in the various systems of a cell, including the interactions between DNA, RNA, proteins and their biosynthesis, as well as the regulation of these interactions. Writing in Nature in 1961, William Astbury described molecular biology as:

...not so much a technique as an approach, an approach from the viewpoint of the so-called basic sciences with the leading idea of searching below the large-scale manifestations of classical biology for the corresponding molecular plan. It is concerned particularly with the forms of biological molecules and [...] is predominantly three-dimensional and structural – which does not mean, however, that it is merely a refinement of morphology. It must at the same time inquire into genesis and function.

Cell biology scientific discipline that studies cells

Cell biology is a branch of biology that studies the structure and function of the cell, which is the basic unit of life. Cell biology is concerned with the physiological properties, metabolic processes, signaling pathways, life cycle, chemical composition and interactions of the cell with their environment. This is done both on a microscopic and molecular level as it encompasses prokaryotic cells and eukaryotic cells. Knowing the components of cells and how cells work is fundamental to all biological sciences; it is also essential for research in bio-medical fields such as cancer, and other diseases. Research in cell biology is closely related to genetics, biochemistry, molecular biology, immunology and cytochemistry.


Illustration from Gray's Anatomy (1918) of a lateral view of the human brain, featuring the hippocampus among other neuroanatomical features Gray739.png
Illustration from Gray's Anatomy (1918) of a lateral view of the human brain, featuring the hippocampus among other neuroanatomical features

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]

Ancient Egypt ancient civilization of Northeastern Africa

Ancient Egypt was a civilization of ancient North Africa, concentrated along the lower reaches of the Nile River in the place that is now the country Egypt. Ancient Egyptian civilization followed prehistoric Egypt and coalesced around 3100 BC with the political unification of Upper and Lower Egypt under Menes. The history of ancient Egypt occurred as a series of stable kingdoms, separated by periods of relative instability known as Intermediate Periods: the Old Kingdom of the Early Bronze Age, the Middle Kingdom of the Middle Bronze Age and the New Kingdom of the Late Bronze Age.

Skull bony structure that forms the skeleton of head in most vertebrates, supports the structures of the face and provides a protective cavity for the brain, composed of two parts: the cranium and the mandible

The skull is a bony structure that forms the head in vertebrates. It supports the structures of the face and provides a protective cavity for the brain. The skull is composed of two parts: the cranium and the mandible. In humans, these two parts are the neurocranium and the viscerocranium or facial skeleton that includes the mandible as its largest bone. The skull forms the anterior most portion of the skeleton and is a product of cephalisation—housing the brain, and several sensory structures such as the eyes, ears, nose, and mouth. In humans these sensory structures are part of the facial skeleton.

The Neolithic, the final division of the Stone Age, began about 12,000 years ago when the first developments of farming appeared in the Epipalaeolithic Near East, and later in other parts of the world. The division lasted until the transitional period of the Chalcolithic from about 6,500 years ago, marked by the development of metallurgy, leading up to the Bronze Age and Iron Age. In Northern Europe, the Neolithic lasted until about 1700 BC, while in China it extended until 1200 BC. Other parts of the world remained broadly in the Neolithic stage of development until European contact.

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 preparation for mummification. It was believed at the time that the heart was the seat of intelligence. According to Herodotus, 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." [10]

Egypt Country spanning North Africa and Southwest Asia

Egypt, officially the Arab Republic of Egypt, is a country spanning the northeast corner of Africa and southwest corner of Asia by a land bridge formed by the Sinai Peninsula. Egypt is a Mediterranean country bordered by the Gaza Strip and Israel to the northeast, the Gulf of Aqaba and the Red Sea to the east, Sudan to the south, and Libya to the west. Across the Gulf of Aqaba lies Jordan, across the Red Sea lies Saudi Arabia, and across the Mediterranean lie Greece, Turkey and Cyprus, although none share a land border with Egypt.

Middle Kingdom of Egypt period in the history of ancient Egypt between about 2000 BC and 1700 BC

The Middle Kingdom of Egypt is the period in the history of ancient Egypt following a period of political division known as the First Intermediate Period. The Middle Kingdom lasted from around 2050 BC to around 1710 BC, stretching from the reunification of Egypt under the reign of Mentuhotep II of the Eleventh Dynasty to the end of the Twelfth Dynasty. The Eleventh Dynasty ruled from Thebes and the Twelfth Dynasty ruled from el-Lisht. Some scholars also include the Thirteenth Dynasty of Egypt wholly into this period as well, in which case the Middle Kingdom would finish around 1650, while others only include it until Merneferre Ay around 1700 BC, last king of this dynasty to be attested in both Upper and Lower Egypt. During the Middle Kingdom period, Osiris became the most important deity in popular religion. The Middle Kingdom was followed by the Second Intermediate Period of Egypt, another period of division that involved foreign invasions of the country by the Hyksos of West Asia.

Mummy Human or animal, whose skin and organs have been preserved

A mummy is a deceased human or an animal whose skin and organs have been preserved by either intentional or accidental exposure to chemicals, extreme cold, very low humidity, or lack of air, so that the recovered body does not decay further if kept in cool and dry conditions. Some authorities restrict the use of the term to bodies deliberately embalmed with chemicals, but the use of the word to cover accidentally desiccated bodies goes back to at least 1615 AD.

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. Plato also speculated that the brain was the seat of the rational part of the soul. [11] Aristotle, however, believed the heart was the center of intelligence and that the brain regulated the amount of heat from the heart. [12] 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.

Ancient Greek medicine

Ancient Greek medicine was a compilation of theories and practices that were constantly expanding through new ideologies and trials. Many components were considered in ancient Greek medicine, intertwining the spiritual with the physical. Specifically, the ancient Greeks believed health was affected by the humors, geographic location, social class, diet, trauma, beliefs, and mindset. Early on the ancient Greeks believed that illnesses were "divine punishments" and that healing was a "gift from the Gods". As trials continued wherein theories were tested against symptoms and results, the pure spiritual beliefs regarding "punishments" and "gifts" were replaced with a foundation based in the physical, i.e., cause and effect.

Hippocrates ancient Greek physician

Hippocrates of Kos, also known as Hippocrates II, was a Greek physician of the Age of Pericles, who is considered one of the most outstanding figures in the history of medicine. He is often referred to as the "Father of Medicine" in recognition of his lasting contributions to the field as the founder of the Hippocratic School of Medicine. This intellectual school revolutionized medicine in ancient Greece, establishing it as a discipline distinct from other fields with which it had traditionally been associated, thus establishing medicine as a profession.

Plato Classical Greek philosopher

Plato was an Athenian philosopher during the Classical period in Ancient Greece, founder of the Platonist school of thought, and the Academy, the first institution of higher learning in the Western world.

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.

Averroes Medieval Arab scholar and philosopher

Ibn Rushd, often Latinized as Averroes, was a Muslim Andalusian philosopher and thinker who wrote about many subjects, including philosophy, theology, medicine, astronomy, physics, Islamic jurisprudence and law, and linguistics. His philosophical works include numerous commentaries on Aristotle, for which he was known in the West as The Commentator. He also served as a judge and a court physician for the Almohad caliphate.

Avicenna medieval Persian polymath, physician, and philosopher

Ibn Sina, also known as Abu Ali Sina, Pur Sina (پورسینا), and often known in the west as Avicenna was a Persian polymath who is regarded as one of the most significant physicians, astronomers, thinkers and writers of the Islamic Golden Age. He has been described as the father of early modern medicine. Of the 450 works he is known to have written, around 240 have survived, including 150 on philosophy and 40 on medicine.

Ibn Zuhr Arab Muslim scholar

Ibn Zuhr, traditionally known by his Latinized name of Avenzoar, was an Arab physician, surgeon, and poet. He was born at Seville in medieval Andalusia, was a contemporary of Averroes and Ibn Tufail, and was the most well-regarded physician of his era. He was particularly known for his emphasis on a more rational, empiric basis of medicine. His major work, Al-Taysīr fil-Mudāwāt wal-Tadbīr, was translated into Latin and Hebrew and was influential to the progress of surgery. He also improved surgical and medical knowledge by keying out several diseases and their treatments.

The Golgi stain first allowed for the visualization of individual neurons. GolgiStainedPyramidalCell.jpg
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 1960, [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 Nervous system diagram-en.svg
Human nervous system

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.

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 is 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 Neuron colored.jpg
Photograph of a stained neuron in a chicken embryo

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. 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 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) Leg Neural Network.jpg
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 Parasagittal MRI of human head in patient with benign familial macrocephaly prior to brain injury (ANIMATED).gif
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.

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.

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

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
Affective neuroscience Affective neuroscience is the study of the neural mechanisms involved in emotion, typically through experimentation on animal models. [40]
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.
Cellular neuroscience Cellular neuroscience is the study of neurons at a cellular level including morphology and physiological properties.
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.
Cultural neuroscience Cultural neuroscience is the study of how cultural values, practices and beliefs shape and are shaped by the mind, brain and genes across multiple timescales. [41]
Developmental neuroscience Developmental neuroscience studies the processes that generate, shape, and reshape the nervous system and seeks to describe the cellular basis of neural development to address underlying mechanisms.
Evolutionary neuroscience Evolutionary neuroscience 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.
Neuroimaging Neuroimaging includes the use of various techniques to either directly or indirectly image the structure and function of the brain.
Neuroimmunology Neuroimmunology is concerned with the interactions between the nervous and the immune system.
Neuroinformatics Neuroinformatics is a discipline within bioinformatics that conducts the organization of neuroscience data and application of computational models and analytical tools.
Neurolinguistics Neurolinguistics is the study of the neural mechanisms in the human brain that control the comprehension, production, and acquisition of language.
Neurophysics Neurophysics deals with the development of physical experimental tools to gain information about the brain.
Neurophysiology Neurophysiology is the study of the functioning of the nervous system, generally using physiological techniques that include measurement and stimulation with electrodes or optically with ion- or voltage-sensitive dyes or light-sensitive channels.
Neuropsychology 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 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.
Paleoneurobiology Paleoneurobiology is a field which combines techniques used in paleontology and archeology to study brain evolution, especially that of the human brain.
Social neuroscience Social neuroscience is an interdisciplinary field devoted to understanding how biological systems implement social processes and behavior, and to using biological concepts and methods to inform and refine theories of social processes and behavior.
Systems neuroscience Systems neuroscience is the study of the function of neural circuits and systems.

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 different countries. [43] 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 . The first National Honor Society in Neuroscience, Nu Rho Psi, was founded in 2006.

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.

YearPrize fieldImageLaureateLifetimeCountryRationaleRef
1904 Physiology Ivan Pavlov nobel.jpg Ivan Petrovich Pavlov 1849–1936Flag of The Russian Empire 1883.svg  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 1843–1926Flag of Italy (1861-1946) crowned.svg  Italy "in recognition of their work on the structure of the nervous system" [56]
Cajal-Restored.jpg Santiago Ramón y Cajal 1852–1934Flag of Spain (1785-1873 and 1875-1931).svg  Spain
1914 Physiology Robert Barany.jpg Robert Bárány 1876–1936Flag of Austria-Hungary (1869-1918).svg  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 1857–1952Flag of the United Kingdom.svg  United Kingdom "for their discoveries regarding the functions of neurons" [58]
Edgar Douglas Adrian nobel.jpg Edgar Douglas Adrian 1889–1977Flag of the United Kingdom.svg  United Kingdom
1936 Physiology Henry Dale nobel.jpg Henry Hallett Dale 1875–1968Flag of the United Kingdom.svg  United Kingdom "for their discoveries relating to chemical transmission of nerve impulses" [59]
Otto Loewi nobel.jpg Otto Loewi 1873–1961Flag of Austria.svg  Austria
Flag of Germany (1935-1945).svg  Germany
1944 Physiology Joseph Erlanger nobel.jpg Joseph Erlanger 1874–1965Flag of the United States (1912-1959).svg  United States "for their discoveries relating to the highly differentiated functions of single nerve fibres" [60]
Herbert Spencer Gasser nobel.jpg Herbert Spencer Gasser 1888–1963Flag of the United States (1912-1959).svg  United States
1949 Physiology Walter Hess.jpg Walter Rudolf Hess 1881–1973Flag of Switzerland.svg   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 1874–1955Flag of Portugal.svg  Portugal "for his discovery of the therapeutic value of leucotomy in certain psychoses" [61]
1957 Physiology Daniel Bovet nobel.jpg Daniel Bovet 1907–1992Flag of Italy.svg  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 Bekesy nobel.jpg Georg von Békésy 1899–1972Flag of the United States.svg  United States "for his discoveries of the physical mechanism of stimulation within the cochlea" [63]
1963 Physiology Eccles lab.jpg John Carew Eccles 1903–1997Flag of Australia (converted).svg  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 1914–1998Flag of the United Kingdom.svg  United Kingdom
Andrew Fielding Huxley nobel.jpg Andrew Fielding Huxley 1917–2012Flag of the United Kingdom.svg  United Kingdom
1967 Physiology Ragnar Granit2.jpg Ragnar Granit 1900–1991Flag of Finland.svg  Finland
Flag of Sweden.svg  Sweden
"for their discoveries concerning the primary physiological and chemical visual processes in the eye" [65]
Haldan Keffer Hartline nobel.jpg Haldan Keffer Hartline 1903–1983Flag of the United States.svg  United States
George Wald nobel.jpg George Wald 1906–1997Flag of the United States.svg  United States
1970 Physiology Julius Axelrod 1912–2004Flag of the United States.svg  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 1905–1983Flag of Sweden.svg  Sweden
Bernard Katz 1911–2003Flag of the United Kingdom.svg  United Kingdom
1981 Physiology Roger Wolcott Sperry.jpg Roger W. Sperry 1913–1994Flag of the United States.svg  United States "for his discoveries concerning the functional specialization of the cerebral hemispheres" [65]
David H. Hubel 1926–2013Flag of Canada (Pantone).svg  Canada "for their discoveries concerning information processing in the visual system" [65]
Torsten Wiesel-7Nov2006.jpg Torsten N. Wiesel 1924-Flag of Sweden.svg  Sweden
1997 Chemistry Skou2008crop.jpg Jens C. Skou 1918–2018Flag of Denmark.svg  Denmark "for the first discovery of an ion-transporting enzyme, Na+, K+ -ATPase" [66]
2000 Physiology Arvid Carlsson 2011a.jpg Arvid Carlsson 1923–2018Flag of Sweden.svg  Sweden "for their discoveries concerning signal transduction in the nervous system" [67]
Paul Greengard.jpg Paul Greengard 1925-Flag of the United States.svg  United States
Eric Richard Kandel.jpg Eric R. Kandel 1929–Flag of the United States.svg  United States
2003 Chemistry Roderick MacKinnon, M.D..jpg Roderick MacKinnon 1956–Flag of the United States.svg  United States "for discoveries concerning channels in cell membranes [...] for structural and mechanistic studies of ion channels" [68]
2014 Physiology John O'Keefe (neuroscientist) 2014b.jpg John O'Keefe 1939–Flag of the United States.svg  United States
Flag of the United Kingdom.svg  United Kingdom
"for their discoveries of cells that constitute a positioning system in the brain" [69]
May-Britt Moser 2014.jpg May-Britt Moser 1963–Flag of Norway.svg  Norway
Edvard Moser.jpg Edvard I. Moser 1962–Flag of Norway.svg  Norway
2017 Physiology Jeffrey C. Hall EM1B8737 (38162359274).jpg Jeffrey C. Hall 1939–Flag of the United States.svg  United States "for their discoveries of molecular mechanisms controlling the circadian rhythm" [70]
Michael Rosbash EM1B8756 (38847326642).jpg Michael Rosbash 1944–Flag of the United States.svg  United States
Michael W. Young D81 4345 (38162439194).jpg Michael W. Young 1949–Flag of the United States.svg  United States

See also

Related Research Articles

Cognitive neuroscience is the scientific field that is concerned with the study of the biological processes and aspects that underlie cognition, with a specific focus on the neural connections in the brain which are involved in mental processes. It addresses the questions of how cognitive activities are affected or controlled by neural circuits in the brain. Cognitive neuroscience is a branch of both neuroscience and psychology, overlapping with disciplines such as behavioral neuroscience, cognitive psychology, physiological psychology and affective neuroscience. Cognitive neuroscience relies upon theories in cognitive science coupled with evidence from neurobiology, and computational modeling.

Gerald Edelman American biologist

Gerald Maurice Edelman was an American biologist who shared the 1972 Nobel Prize in Physiology or Medicine for work with Rodney Robert Porter on the immune system. Edelman's Nobel Prize-winning research concerned discovery of the structure of antibody molecules. In interviews, he has said that the way the components of the immune system evolve over the life of the individual is analogous to the way the components of the brain evolve in a lifetime. There is a continuity in this way between his work on the immune system, for which he won the Nobel Prize, and his later work in neuroscience and in philosophy of mind.

Eric Kandel American neuropsychiatrist

Eric Richard Kandel is an Austrian-American medical doctor who specialized in Psychiatry, a neuroscientist and a University Professor of biochemistry and biophysics at the College of Physicians and Surgeons at Columbia University. He was a recipient of the 2000 Nobel Prize in Physiology or Medicine for his research on the physiological basis of memory storage in neurons. He shared the prize with Arvid Carlsson and Paul Greengard.

The following outline is provided as an overview of and topical guide to neuroscience:

Behavioral neuroscience, also known as biological psychology, biopsychology, or psychobiology, is the application of the principles of biology to the study of physiological, genetic, and developmental mechanisms of behavior in humans and other animals.

Systems neuroscience is a subdiscipline of neuroscience and systems biology that studies the structure and function of neural circuits and systems. It is an umbrella term, encompassing a number of areas of study concerned with how nerve cells behave when connected together to form neural pathways, neural circuits, and larger brain networks. At this level of analysis, neuroscientists study how different neural circuits analyze sensory information, form perceptions of the external world, make decisions, and execute movements. Researchers in systems neuroscience are concerned with the relation between molecular and cellular approaches to understanding brain structure and function, as well as with the study of high-level mental functions such as language, memory, and self-awareness. Systems neuroscientists typically employ techniques for understanding networks of neurons as they are seen to function, by way of electrophysiology using either single-unit recording or multi-electrode recording, functional magnetic resonance imaging (fMRI), and PET scans. The term is commonly used in an educational framework: a common sequence of graduate school neuroscience courses consists of cellular/molecular neuroscience for the first semester, then systems neuroscience for the second semester. It is also sometimes used to distinguish a subdivision within a neuroscience department at an academic institution.

Paul Greengard American neuroscientist

Paul Greengard was an American neuroscientist best known for his work on the molecular and cellular function of neurons. In 2000, Greengard, Arvid Carlsson and Eric Kandel were awarded the Nobel Prize for Physiology or Medicine for their discoveries concerning signal transduction in the nervous system. He was Vincent Astor Professor at Rockefeller University, and served on the Scientific Advisory Board of the Cure Alzheimer's Fund, as well as the Scientific Council of the Brain & Behavior Research Foundation. He was married to artist Ursula von Rydingsvard.

Terrence (Terry) Joseph Sejnowski is the Francis Crick Professor at the Salk Institute for Biological Studies where he directs the Computational Neurobiology Laboratory and is the director of the Crick-Jacobs center for theoretical and computational biology. His research in neural networks and computational neuroscience has been pioneering.

Neuroinformatics is a research field concerned with the organization of neuroscience data by the application of computational models and analytical tools. These areas of research are important for the integration and analysis of increasingly large-volume, high-dimensional, and fine-grain experimental data. Neuroinformaticians provide computational tools, mathematical models, and create interoperable databases for clinicians and research scientists. Neuroscience is a heterogeneous field, consisting of many and various sub-disciplines. In order for our understanding of the brain to continue to deepen, it is necessary that these sub-disciplines are able to share data and findings in a meaningful way; Neuroinformaticians facilitate this.

<i>Principles of Neural Science</i>

First published in 1981 by Elsevier, Principles of Neural Science is an influential neuroscience textbook edited by Eric R. Kandel, James H. Schwartz, and Thomas M. Jessell. The original edition was 468 pages; now on the fifth edition, the book has grown to 1747 pages. The second edition was published in 1985, third in 1991, fourth in 2000. The fifth and latest edition was published on October 26th, 2012 and includes Steven A. Siegelbaum and A.J. Hudspeth as editors. It has been hailed as the "Bible of Neuroscience" having been co-written by the "Father of Neuroscience", Eric Kandel.

From the ancient Egyptian mummifications to 18th century scientific research on "globules" and neurons, there is evidence of neuroscience practice throughout the early periods of history. The early civilizations lacked adequate means to obtain knowledge about the human brain. Their assumptions about the inner workings of the mind, therefore, were not accurate. Early views on the function of the brain regarded it to be a form of "cranial stuffing" of sorts. In ancient Egypt, from the late Middle Kingdom onwards, in preparation for mummification, the brain was regularly removed, for it was the heart that was assumed to be the seat of intelligence. According to Herodotus, during the first step of mummification: "The most perfect practice is to extract as much of the brain as possible with an iron hook, and what the hook cannot reach is mixed with drugs." Over the next five thousand years, this view came to be reversed; the brain is now known to be the seat of intelligence, although colloquial variations of the former remain as in "memorizing something by heart".

Sten Grillner is a Swedish neurophysiologist and distinguished professor at the Karolinska Institute's Nobel Institute for Neurophysiology in Stockholm where he is the director of the institute. He is considered one of the world’s foremost experts in the cellular bases of motor behaviour. His research is focused on understanding the cellular bases of motor behaviour; in particular, he has shown how neuronal circuits in the spine help control rhythmic movements, such as those needed for locomotion. He is current secretary general of International Brain Research Organization IBRO and President of the Federation of European Neuroscience Societies (FENS). For his work, in 2008 he was awarded the $1 million Kavli Prize for deciphering the basic mechanisms which govern the development and functioning of the networks of cells in the brain and spinal cord. This prize distinguish the recipient from the Nobel prizes in basic medical sciences.

Eve Marder American neuroscientist

Eve Marder is a University Professor and the Victor and Gwendolyn Beinfield Professor of Neuroscience at Brandeis University. At Brandeis, Marder is also a member of the Volen National Center for Complex Systems. Dr. Marder is known for her pioneering work on small neuronal networks which her team has interrogated via a combination of complementary experimental and theoretical techniques.

Some of the research that is conducted in the field of psychology is more "fundamental" than the research conducted in the applied psychological disciplines, and does not necessarily have a direct application. The subdisciplines within psychology that can be thought to reflect a basic-science orientation include biological psychology, cognitive psychology, neuropsychology, and so on. Research in these subdisciplines is characterized by methodological rigor. The concern of psychology as a basic science is in understanding the laws and processes that underlie behavior, cognition, and emotion. Psychology as a basic science provides a foundation for applied psychology. Applied psychology, by contrast, involves the application of psychological principles and theories yielded up by the basic psychological sciences; these applications are aimed at overcoming problems or promoting well-being in areas such as mental and physical health and education.

Consciousness after death

Consciousness after death is a common theme in society and culture in the context of life after death. Scientific research has established that the mind and consciousness are closely connected with the physiological functioning of the brain, the cessation of which defines brain death. However, many believe in some form of life after death, which is a feature of many religions.

John OKeefe (neuroscientist) American British neuroscientist, 2014 Nobel laureate in Physiology or Medicine

John O'Keefe, is an American-British neuroscientist and a professor at the Sainsbury Wellcome Centre for Neural Circuits and Behaviour and the Research Department of Cell and Developmental Biology at University College London. He discovered place cells in the hippocampus, and that they show a specific kind of temporal coding in the form of theta phase precession. In 2014 he received the Kavli Prize in Neuroscience "for the discovery of specialized brain networks for memory and cognition", together with Brenda Milner and Marcus Raichle. He shared the Nobel Prize in Physiology or Medicine also that year, together with May-Britt Moser and Edvard Moser.

The Karl Spencer Lashley Award is awarded by The American Philosophical Society as a recognition of research on the integrative neuroscience of behavior. The award was established in 1957 by a gift from Dr. Karl Spencer Lashley.

The Princeton Neuroscience Institute (PNI) is a center for neuroscience research at Princeton University. Founded in the spring of 2004, the PNI serves as a "stimulus for teaching and research in neuroscience and related fields" and "places particular emphasis on the close connection between theory, modeling, and experimentation using the most advanced technologies." It often partners with Princeton University's departments of Psychology and Molecular Biology.


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Further reading