The scope of neuroscience has broadened over time to include different approaches used to study the nervous system at different scales. The techniques used byneuroscientists have expanded enormously, from molecular andcellular studies of individual neurons toimaging ofsensory,motor, andcognitive tasks in the brain.
The earliest study of the nervous system dates toancient Egypt.Trepanation, the surgical practice of either drilling or scraping a hole into theskull for the purpose of curing head injuries ormental disorders, or relieving cranial pressure, was first recorded during theNeolithic period. Manuscripts dating to1700 BC indicate that theEgyptians had some knowledge about symptoms ofbrain damage.[10]
Early views on the function of the brain regarded it to be a "cranial stuffing" of sorts. InEgypt, from the lateMiddle Kingdom onwards, the brain was regularly removed in preparation formummification. It was believed at the time that theheart was the seat of intelligence. According toHerodotus, 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 theskull is cleared of the rest by rinsing with drugs."[11]
The view that the heart was the source of consciousness was not challenged until the time of theGreek physicianHippocrates. He believed that the brain was not only involved with sensation—since most specialized organs (e.g., eyes, ears, tongue) are located in the head near the brain—but was also the seat of intelligence.[12]Plato also speculated that the brain was the seat of the rational part of the soul.[13]Aristotle, however, believed the heart was the center of intelligence and that the brain regulated the amount of heat from the heart.[14] This view was generally accepted until theRoman physicianGalen, a follower of Hippocrates and physician toRoman gladiators, observed that his patients lost their mental faculties when they had sustained damage to their brains.[15]
TheGolgi stain first allowed for the visualization of individual neurons.
Luigi Galvani's pioneering work in the late 1700s set the stage for studying theelectrical excitability of muscles and neurons. In 1843Emil du Bois-Reymond demonstrated the electrical nature of the nerve signal,[16] whose speedHermann von Helmholtz proceeded to measure,[17] and in 1875Richard Caton found electrical phenomena in the cerebral hemispheres of rabbits and monkeys.[18]Adolf Beck published in 1890 similar observations of spontaneous electrical activity of the brain of rabbits and dogs.[19] Studies of the brain became more sophisticated after the invention of themicroscope and the development of astaining procedure byCamillo Golgi during the late 1890s. The procedure used asilver chromate salt to reveal the intricate structures of individualneurons. His technique was used bySantiago Ramón y Cajal and led to the formation of theneuron doctrine, the hypothesis that the functional unit of the brain is the neuron.[20] Golgi and Ramón y Cajal shared theNobel 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 1815Jean 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 byMarc Dax in 1836 andPaul Broca in 1865 suggested that certain regions of the brain were responsible for certain functions.[21] At the time, these findings were seen as a confirmation ofFranz Joseph Gall's theory that language was localized and that certainpsychological functions were localized in specific areas of thecerebral cortex.[22][23] Thelocalization of function hypothesis was supported by observations ofepileptic patients conducted byJohn Hughlings Jackson, who correctly inferred the organization of themotor 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. In 1894, neurologist and psychiatristEdward Flatau published a human brain atlas “Atlas of the Human Brain and the Course of the Nerve-Fibres” which consisted of long-exposure photographs of fresh brain sections.[24] In 1897,Charles Scott Sherrington introduced the name "synapse" for the connection between neurons.[25]
Brodmann's diagram of the cerebral cortex with the areas he identified
In 1909, German anatomistKorbinian Brodmann published his original research on brain mapping, defining 52 distinct regions of the cerebral cortex, known asBrodmann areas.[26] Modern research throughneuroimaging techniques, still uses theBrodmanncerebral cytoarchitectonic map (referring to the study ofcell structure) anatomical definitions from this era in continuing to show that distinct areas of the cortex are activated in the execution of specific tasks.[27]
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 citedDavid Rioch,Francis O. Schmitt, andStephen Kuffler as having played critical roles in establishing the field.[28] Rioch originated the integration of basic anatomical and physiological research with clinical psychiatry at theWalter Reed Army Institute of Research, starting in the 1950s. During the same period, Schmitt established a neuroscience research program within the Biology Department at theMassachusetts 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 byJames L. McGaugh.[29] This was followed by theDepartment of Neurobiology atHarvard Medical School, which was founded in 1966 by Stephen Kuffler.[30]
In the process of treatingepilepsy,Wilder Penfield produced maps of the location of various functions (motor, sensory, memory, vision) in the brain.[31][32] He summarized his findings in a 1950 book calledThe Cerebral Cortex of Man.[33] Wilder Penfield and his co-investigators Edwin Boldrey and Theodore Rasmussen are considered to be the originators of thecortical homunculus.[34]
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 andAndrew Huxley presented amathematical model for the transmission of electrical signals in neurons of the giant axon of a squid, which they called "action potentials", and how they are initiated and propagated, known as theHodgkin–Huxley model. In 1961–1962, Richard FitzHugh and J. Nagumo simplified Hodgkin–Huxley, in what is called theFitzHugh–Nagumo model. In 1962,Bernard Katz modeledneurotransmission across the space between neurons known assynapses. Beginning in 1966, Eric Kandel and collaborators examined biochemical changes in neurons associated with learning and memory storage inAplysia. In 1981 Catherine Morris and Harold Lecar combined these models in theMorris–Lecar model. Such increasingly quantitative work gave rise to numerousbiological neuron models andmodels of neural computation.
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.[43]
Thescientific study of the nervous system increased significantly during the second half of the twentieth century, principally due to advances inmolecular biology,electrophysiology, andcomputational neuroscience. This has allowed neuroscientists to study thenervous 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 singleneuron. Neurons are cells specialized for communication. They are able to communicate with neurons and other cell types through specialized junctions calledsynapses, at which electrical or electrochemical signals can be transmitted from one cell to another. Many neurons extrude a long thin filament ofaxoplasm called anaxon, 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 nervoussystem emerges from the assemblage of neurons that are connected to each other inneural circuits, andnetworks.
The vertebrate nervous system can be split into two parts: thecentral nervous system (defined as thebrain andspinal cord), and theperipheral nervous system. In many species—including all vertebrates—the nervous system is the mostcomplex organ system in the body, with most of the complexity residing in the brain. Thehuman 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.[44]
Due to the high degree ofplasticity of the human brain, the structure of its synapses and their resulting functions change throughout life.[45]
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 inelectron microscopy,computer science,electronics,functional neuroimaging, andgenetics andgenomics have all been major drivers of progress.
Advances in the classification ofbrain cells have been enabled by electrophysiological recording,single-cell genetic sequencing, and high-quality microscopy, which have combined into a single method pipeline calledpatch-sequencing in which all three methods are simultaneously applied using miniature tools.[46] 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.[47]
Basic questions addressed inmolecular neuroscience include the mechanisms by which neurons express and respond to molecular signals and howaxons form complex connectivity patterns. At this level, tools frommolecular biology andgenetics are used to understand how neurons develop and how genetic changes affect biological functions.[48] Themorphology, molecular identity, and physiological characteristics of neurons and how they relate to different types of behavior are also of considerable interest.[49]
Questions addressed incellular neuroscience include the mechanisms of how neurons processsignals physiologically and electrochemically. These questions include how signals are processed by neurites and somas and howneurotransmitters and electrical signals are used to process information in a neuron. Neurites are thin extensions from a neuronalcell body, consisting ofdendrites (specialized to receive synaptic inputs from other neurons) andaxons (specialized to conduct nerve impulses calledaction potentials). Somas are the cell bodies of the neurons and contain the nucleus.[50]
Computational neurogenetic modeling is concerned with the development of dynamic neuronal models for modeling brain functions with respect to genes and dynamic interactions between genes, on the cellular level (Computational Neurogenetic Modeling (CNGM) can also be used to model neural systems).[55]
Proposed organization of motor-semantic neural circuits for action language comprehension. Adapted from Shebani et al. (2013).
Systems neuroscience research centers on the structural and functional architecture of the developing human brain, and the functions oflarge-scale brain networks, or functionally-connected systems within the brain. Alongside brain development, systems neuroscience also focuses on how the structure and function of the brain enables or restricts the processing of sensory information, using learnedmental models of the world, to motivate behavior.
Questions in systems neuroscience include howneural circuits are formed and used anatomically and physiologically to produce functions such asreflexes,multisensory integration,motor coordination,circadian rhythms,emotional responses,learning, andmemory.[56] 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.[57]
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.[58]
Cognitive neuroscience addresses the questions of howpsychological functions are produced byneural circuitry. The emergence of powerful new measurement techniques such asneuroimaging (e.g.,fMRI,PET,SPECT),EEG,MEG,electrophysiology,optogenetics andhuman genetic analysis combined with sophisticatedexperimental techniques fromcognitive psychology allowsneuroscientists andpsychologists 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.[62] Moreover, the neuroscientific identification of multiple memory systems related to different brain areas has challenged the idea ofmemory as a literal reproduction of the past, supporting a view of memory as a generative, constructive and dynamic process.[63]
Questions in computational neuroscience can span a wide range of levels of traditional analysis, such asdevelopment,structure, andcognitive functions of the brain. Research in this field utilizesmathematical models, theoretical analysis, andcomputer 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 ofneural networks. Computational neuroscience is often referred to as theoretical neuroscience.
Neurology works with diseases of the central and peripheral nervous systems, such asamyotrophic lateral sclerosis (ALS) andstroke, and their medical treatment.Psychiatry focuses onaffective, behavioral,cognitive, andperceptual 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 andpsychosurgery work primarily with surgical treatment of diseases of the central and peripheral nervous systems.[66]
Neuroscience underlies the development of variousneurotherapy methods to treat diseases of the nervous system.[67][68][69]
Recently, the boundaries between various specialties have blurred, as they are all influenced bybasic research in neuroscience. For example,brain imaging enables objective biological insight into mental illnesses, which can lead to faster diagnosis, more accurate prognosis, and improved monitoring of patient progress over time.[70]
Integrative neuroscience describes the effort to combine models and information from multiple levels of research to develop a coherent model of the nervous system. For example, brain imaging coupled with physiological numerical models and theories of fundamental mechanisms may shed light on psychiatric disorders.[71]
Another important area of translational research isbrain–computer interfaces (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.[72] Translational BCI research is supported by specialized neurotechnology platforms that enable high-resolution neural signal acquisition, real-time processing, and experimental validation in clinical and laboratory settings. Such platforms are developed by academic groups as well as industry partners, including systems produced by g.tec medical engineering GmbH.[73]
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.
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.[75]
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.[79]
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.[80]
Neuroinformatics is a discipline within bioinformatics that conducts the organization of neuroscience data and application of computational models and analytical tools.[91]
Neurolinguistics is the study of the neural mechanisms in the human brain that control the comprehension, production, and acquisition of language.[92][77]
Neuro-ophthalmology is an academically oriented subspecialty that merges the fields of neurology and ophthalmology, often dealing with complex systemic diseases that have manifestations in the visual system.
Neurophysiology is the study of the structure and function 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.[94]
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 withbiopsychology,clinical psychology,cognitive psychology, anddevelopmental 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.[95]
Neuropsychopharmacology, an interdisciplinary science related topsychopharmacology and fundamental neuroscience, is the study of the neural mechanisms that drugs act upon to influence behavior.[96]
Paleoneurobiology is a field that combines techniques used in paleontology and archeology to study brain evolution, especially that of the human brain.[97]
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.[98]
The career options for neuroscience graduates vary widely depending on the level of education. At the bachelor’s level, graduates often enter laboratory research, healthcare support, biotechnology, or science communication, though some pursue broader fields such as policy or nonprofit work. With a master’s degree, training may prepare individuals for applied health professions (e.g., occupational therapy, medicine -neurology, psychiatry, neuroimaging-, genetic counseling), research management, or public health. An advanced degree (PhD or equivalent) is usually required for independent research or university teaching.Source:[100]
The largest professional neuroscience organization is theSociety for Neuroscience (SFN), which is based in theUnited States but includes many members from other countries. Since its founding in 1969 the SFN has grown steadily: as of 2010 it recorded 40,290 members from 83 countries.[101] 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 theInternational Brain Research Organization (IBRO), which holds its meetings in a country from a different part of the world each year, and theFederation 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 theBritish Neuroscience Association, the German Neuroscience Society (Neurowissenschaftliche Gesellschaft), and the FrenchSociété des Neurosciences.[102] 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[103] and Project Encephalon.[104]
In 2013, theBRAIN Initiative was announced in the US. The International Brain Initiative[105] was created in 2017,[106] currently integrated by more than seven national-level brain research initiatives (US,Europe,Allen Institute,Japan,China, Australia,[107] Canada,[108] Korea,[109] and Israel[110])[111] spanning four continents.
In addition to conducting traditional research in laboratory settings, neuroscientists have also been involved in thepromotion 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 theInternational Brain Bee, which is an academic competition for high school or secondary school students worldwide.[112] In the United States, large organizations such as the Society for Neuroscience have promoted neuroscience education by developing a primer called Brain Facts,[113] collaborating with public school teachers to develop Neuroscience Core Concepts for K-12 teachers and students,[114] and cosponsoring a campaign with theDana Foundation called Brain Awareness Week to increase public awareness about the progress and benefits of brain research.[115] In Canada, the Canadian Institutes of Health Research's (CIHR) Canadian National Brain Bee is held annually atMcMaster University.[116]
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.[117]
Neuroscientists have also collaborated with other education experts to study and refine educational techniques to optimize learning among students, an emerging field callededucational neuroscience.[118] Federal agencies in the United States, such as theNational Institute of Health (NIH)[119] andNational Science Foundation (NSF),[120] have also funded research that pertains to best practices in teaching and learning of neuroscience concepts.
Neuromorphic engineering is a branch of neuroscience that deals with creating functionalphysical models of neurons for the purposes of useful computation.[121][122] The emergent computational properties of neuromorphic computers are fundamentally different from conventional computers in the sense that they arecomplex systems, and that the computational components are interrelated with no central processor.[123]
One example of such a computer is theSpiNNaker supercomputer.[124]
Sensors can also be made smart with neuromorphic technology. An example of this is theEvent 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 theHuman 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.[125]
Recent advances inneuromorphic microchip technology have led a group of scientists to create an artificial neuron that can replace real neurons in diseases.[126][127]
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