Array ( [0] => {{Short description|Ability of the brain to continuously change}} [1] => {{cs1 config|name-list-style=vanc|display-authors=6}} [2] => {{Redirect|Neural plasticity|the journal|Neural Plasticity (journal)|the 2014 Cold Specks album|Neuroplasticity (album)}} [3] => {{Use dmy dates|date= July 2015}} [4] => [5] => '''Neuroplasticity''', also known as '''neural plasticity''' or '''brain plasticity''', is the ability of [[neural network]]s in the [[brain]] to change through [[neurogenesis|growth]] and reorganization. It is when the brain is rewired to function in some way that differs from how it previously functioned.{{Cite book |author=Costandi, Moheb |url=http://worldcat.org/oclc/987683015 |title=Neuroplasticity |date=19 August 2016 |publisher=MIT Press |isbn=978-0-262-52933-4 |oclc=987683015}} These changes range from individual [[neuron pathways]] making new connections, to systematic adjustments like [[cortical remapping]] or [[neural oscillation]]. Other forms of neuroplasticity include homologous area adaptation, cross modal reassignment, map expansion, and compensatory masquerade.{{Cite journal |last=Grafman |first=Jordan |date=July 1, 2000 |title=Conceptualizing functional neuroplasticity |url=https://linkinghub.elsevier.com/retrieve/pii/S0021992400000307 |journal=Journal of Communication Disorders |language=en |volume=33 |issue=4 |pages=345–356 |doi=10.1016/S0021-9924(00)00030-7|pmid=11001161 }} Examples of neuroplasticity include [[neural circuit|circuit]] and network changes that result from [[learning]] a new ability, [[neural coding|information acquisition]], environmental influences, pregnancy,{{Cite journal |last1=Paternina-Die |first1=María |last2=Martínez-García |first2=Magdalena |last3=Martín de Blas |first3=Daniel |last4=Noguero |first4=Inés |last5=Servin-Barthet |first5=Camila |last6=Pretus |first6=Clara |last7=Soler |first7=Anna |last8=López-Montoya |first8=Gonzalo |last9=Desco |first9=Manuel |last10=Carmona |first10=Susana |date=February 2024 |title=Women's neuroplasticity during gestation, childbirth and postpartum |journal=Nature Neuroscience |language=en |volume=27 |issue=2 |pages=319–327 |doi=10.1038/s41593-023-01513-2 |pmid=38182834 |pmc=10849958 |issn=1546-1726}} caloric intake, practice/training, and [[psychological stress]]. [6] => [7] => Neuroplasticity was once thought by [[neuroscientist]]s to manifest only during childhood,{{cite journal | vauthors = Leuner B, Gould E | title = Structural plasticity and hippocampal function | journal = Annual Review of Psychology | volume = 61 | issue = 1 | pages = 111–140 | date = January 2010 | pmid = 19575621 | pmc = 3012424 | doi = 10.1146/annurev.psych.093008.100359 }}{{Cite book| vauthors = Kusiak AN, Selzer ME |chapter-url= https://books.google.com/books?id=Vb9zDAAAQBAJ&pg=PT6|title= Neurological Rehabilitation |date= 2013|publisher= Elsevier Inc. Chapters|isbn= 978-0-12-807792-4| veditors = Barnes MP, Good DC | edition = 3rd |location= China|language= en|chapter= Neuroplasticity in the spinal cord|access-date= 3 June 2020|archive-date= 13 July 2020|archive-url= https://web.archive.org/web/20200713022512/https://books.google.com/books?id=Vb9zDAAAQBAJ&lpg=PT6|url-status= live}} but research in the latter half of the 20th century showed that many aspects of the brain can be altered (or are "plastic") even through adulthood. However, the developing brain exhibits a higher degree of plasticity than the adult brain.{{cite journal | vauthors = Hensch TK, Bilimoria PM | title = Re-opening Windows: Manipulating Critical Periods for Brain Development | journal = Cerebrum | volume = 2012 | pages = 11 | date = July 2012 | pmid = 23447797 | pmc = 3574806 }} [[Activity-dependent plasticity]] can have significant implications for healthy development, learning, [[memory]], and recovery from [[brain damage]].{{cite journal | vauthors = Ganguly K, Poo MM | title = Activity-dependent neural plasticity from bench to bedside | journal = Neuron | volume = 80 | issue = 3 | pages = 729–741 | date = October 2013 | pmid = 24183023 | doi = 10.1016/j.neuron.2013.10.028 | doi-access = free }}{{cite journal | vauthors = Carey L, Walsh A, Adikari A, Goodin P, Alahakoon D, De Silva D, Ong KL, Nilsson M, Boyd L | title = Finding the Intersection of Neuroplasticity, Stroke Recovery, and Learning: Scope and Contributions to Stroke Rehabilitation | journal = Neural Plasticity | volume = 2019 | pages = 5232374 | date = 2 May 2019 | pmid = 31191637 | pmc = 6525913 | doi = 10.1155/2019/5232374 | doi-access = free }} [8] => [9] => == History == [10] => ===Origin=== [11] => The term ''plasticity'' was first applied to behavior in 1890 by [[William James]] in ''[[The Principles of Psychology]]'' where the term was used to describe "a structure weak enough to yield to an influence, but strong enough not to yield all at once".{{Cite journal |last1=Warraich |first1=Zuha |last2=Kleim |first2=Jeffrey A. |date=2010-12-01 |title=Neural Plasticity: The Biological Substrate For Neurorehabilitation |url=http://doi.wiley.com/10.1016/j.pmrj.2010.10.016 |journal=PM&R |language=en |volume=2 |issue=12 Suppl 2 |pages=S208–S219 |doi=10.1016/j.pmrj.2010.10.016|pmid=21172683 |s2cid=36928880 }} The first person to use the term ''neural plasticity'' appears to have been the Polish neuroscientist [[Jerzy Konorski]]. [12] => [13] => One of the first experiments providing evidence for neuroplasticity was conducted in 1793, by Italian anatomist Michele Vicenzo Malacarne, who described experiments in which he paired animals, trained one of the pair extensively for years, and then dissected both. Malacarne discovered that the cerebellums of the trained animals were substantially larger than the cerebellum of the untrained animals. However, while these findings were significant, they were eventually forgotten.{{cite journal | vauthors = Rosenzweig MR | title = Aspects of the search for neural mechanisms of memory | journal = Annual Review of Psychology | volume = 47 | pages = 1–32 | year = 1996 | pmid = 8624134 | doi = 10.1146/annurev.psych.47.1.1 | doi-access = }} In 1890, the idea that the brain and its function are not fixed throughout adulthood was proposed by [[William James]] in ''[[The Principles of Psychology]]'', though the idea was largely neglected. Up until the 1970s, neuroscientists believed that the brain's structure and function was essentially fixed throughout adulthood. [14] => [15] => While the brain was commonly understood as a nonrenewable organ in the early 1900s, [[Santiago Ramón y Cajal]], ''father of neuroscience'', used the term neuronal plasticity to describe nonpathological changes in the structure of adult brains. Based on his renowned [[neuron doctrine]], Cajal first described the neuron as the fundamental unit of the nervous system that later served as an essential foundation to develop the concept of neural plasticity. Many neuroscientists used the term plasticity to explain the regenerative capacity of the [[peripheral nervous system]] only. Cajal, however, used the term plasticity to reference his findings of degeneration and regeneration in the adult brain (a part of the [[central nervous system]]). This was controversial.{{cite journal | vauthors = Fuchs E, Flügge G | title = Adult neuroplasticity: more than 40 years of research | journal = Neural Plasticity | volume = 2014 | issue = 5 | pages = 541870 | date = 2014 | pmid = 24883212 | pmc = 4026979 | doi = 10.1155/2014/541870 | doi-access = free }} [16] => [17] => The term has since been broadly applied: [18] => {{blockquote|Given the central importance of neuroplasticity, an outsider would be forgiven for assuming that it was well defined and that a basic and universal framework served to direct current and future hypotheses and experimentation. Sadly, however, this is not the case. While many neuroscientists use the word neuroplasticity as an umbrella term it means different things to different researchers in different subfields ... In brief, a mutually agreed-upon framework does not appear to exist. [19] => }} [20] => [21] => === Research and discovery === [22] => [23] => In 1923, [[Karl Lashley]] conducted experiments on [[rhesus monkey]]s that demonstrated changes in neuronal pathways, which he concluded were evidence of plasticity. Despite this, and other research that suggested plasticity, neuroscientists did not widely accept the idea of neuroplasticity. [24] => [25] => In 1945, [[Justo Gonzalo]] concluded from his research on brain dynamics, that, contrary to the activity of the [[projection areas]], the "central" cortical mass (more or less equidistant from the visual, tactile and auditive projection areas), would be a "maneuvering mass", rather unspecific or multisensory, with capacity to increase neural excitability and re-organize the activity by means of plasticity properties. He gives as a first example of adaptation, to see upright with reversing glasses in the [[George M. Stratton|Stratton]] experiment,{{cite journal | vauthors = Stratton GM | s2cid = 13147419 | author-link = George M. Stratton | year = 1896 | title = Some preliminary experiments on vision without inversion of the retinal image | journal = Psychological Review | volume = 3 | issue = 6| pages = 611–7 | doi = 10.1037/h0072918}} and specially, several first-hand brain injuries cases in which he observed dynamic and adaptive properties in their disorders, in particular in the inverted perception disorder [e.g., see pp 260–62 Vol. I (1945), p 696 Vol. II (1950)]. He stated that a sensory signal in a projection area would be only an inverted and constricted outline that would be magnified due to the increase in recruited cerebral mass, and re-inverted due to some effect of brain plasticity, in more central areas, following a spiral growth.{{Cite journal [26] => | author-link = Justo Gonzalo [27] => |journal = Trabajos del Instituto Cajal de Investigaciones Biológicas [28] => |pages=95–157 | volume=44 | year=1952 [29] => |last=Gonzalo|first=Justo|access-date=2012-04-12|title=Dinámica cerebral|hdl = 10347/4341 [30] => |url=http://hdl.handle.net/10347/4341 [31] => }} [32] => [33] => [[Marian Diamond]] of the University of California, Berkeley, produced the first scientific evidence of anatomical brain plasticity, publishing her research in 1964.{{cite journal | vauthors = Diamond MC, Krech D, Rosenzweig MR | title = The effects of an enriched environment on the histology of the rat cerebral cortex | journal = The Journal of Comparative Neurology | volume = 123 | pages = 111–120 | date = August 1964 | pmid = 14199261 | doi = 10.1002/cne.901230110 | s2cid = 30997263 }}{{cite journal | vauthors = Bennett EL, Diamond MC, Krech D, Rosenzweig MR | title = Chemical and Anatomical Plasticity of Brain | journal = Science | volume = 146 | issue = 3644 | pages = 610–619 | date = October 1964 | pmid = 14191699 | doi = 10.1126/science.146.3644.610 | bibcode = 1964Sci...146..610B }} [34] => [35] => Other significant evidence was produced in the 1960s and after, notably from scientists including [[Paul Bach-y-Rita]], [[Michael Merzenich]] along with [[Jon Kaas]], as well as several others.''Brain Science Podcast'' Episode #10, "Neuroplasticity" [36] => [37] => In the 1960s, [[Paul Bach-y-Rita]] invented a device that was tested on a small number of people, and involved a person sitting in a chair, embedded in which were nubs that were made to vibrate in ways that translated images received in a camera, allowing a form of vision via [[sensory substitution]].{{cite web |url=https://www.pbs.org/kcet/wiredscience/video/286-mixed_feelings.html |title=Wired Science . Video: Mixed Feelings |publisher=PBS |access-date=12 June 2010 |archive-date=22 December 2007 |archive-url=https://web.archive.org/web/20071222071618/https://www.pbs.org/kcet/wiredscience/video/286-mixed_feelings.html |url-status=live }} [38] => [39] => Studies in people recovering from [[stroke]] also provided support for neuroplasticity, as regions of the brain that remained healthy could sometimes take over, at least in part, functions that had been destroyed; [[Shepherd Ivory Franz]] did work in this area.{{cite web|url=http://rkthomas.myweb.uga.edu/Franz.htm|archive-url=https://web.archive.org/web/20120203003746/http://rkthomas.myweb.uga.edu/Franz.htm|archive-date=2012-02-03 |title=Shepherd Ivory Franz |publisher=Rkthomas.myweb.uga.edu |access-date=12 June 2010}}{{cite journal | vauthors = Colotla VA, Bach-y-Rita P | title = Shepherd Ivory Franz: his contributions to neuropsychology and rehabilitation | journal = Cognitive, Affective & Behavioral Neuroscience | volume = 2 | issue = 2 | pages = 141–148 | date = June 2002 | pmid = 12455681 | doi = 10.3758/CABN.2.2.141 | url = http://htpprints.yorku.ca/archive/00000236/01/Colotla_Bach-y-Rita_2002.pdf | url-status = unfit | s2cid = 45175011 | doi-access = free | archive-url = https://web.archive.org/web/20120301092815/http://htpprints.yorku.ca/archive/00000236/01/Colotla_Bach-y-Rita_2002.pdf | archive-date = 1 March 2012 }} [40] => [41] => [[Eleanor Maguire]] documented changes in hippocampal structure associated with acquiring the knowledge of London's layout in local taxi drivers.{{cite journal | vauthors = Maguire EA, Frackowiak RS, Frith CD | title = Recalling routes around london: activation of the right hippocampus in taxi drivers | journal = The Journal of Neuroscience | volume = 17 | issue = 18 | pages = 7103–7110 | date = September 1997 | pmid = 9278544 | pmc = 6573257 | doi = 10.1523/JNEUROSCI.17-18-07103.1997 }}{{cite journal | vauthors = Woollett K, Maguire EA | title = Acquiring "the Knowledge" of London's layout drives structural brain changes | journal = Current Biology | volume = 21 | issue = 24 | pages = 2109–2114 | date = December 2011 | pmid = 22169537 | pmc = 3268356 | doi = 10.1016/j.cub.2011.11.018 | bibcode = 2011CBio...21.2109W }}{{cite journal | vauthors = Maguire EA, Gadian DG, Johnsrude IS, Good CD, Ashburner J, Frackowiak RS, Frith CD | title = Navigation-related structural change in the hippocampi of taxi drivers | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 97 | issue = 8 | pages = 4398–4403 | date = April 2000 | pmid = 10716738 | pmc = 18253 | doi = 10.1073/pnas.070039597 | doi-access = free | bibcode = 2000PNAS...97.4398M }} A redistribution of grey matter was indicated in London Taxi Drivers compared to controls. This work on hippocampal plasticity not only interested scientists, but also engaged the public and media worldwide. [42] => [43] => [[Michael Merzenich]] is a neuroscientist who has been one of the pioneers of neuroplasticity for over three decades. He has made some of "the most ambitious claims for the field – that brain exercises may be as useful as drugs to treat diseases as severe as schizophrenia – that plasticity exists from cradle to the grave, and that radical improvements in cognitive functioning – how we learn, think, perceive, and remember are possible even in the elderly." Merzenich's work was affected by a crucial discovery made by [[David Hubel]] and [[Torsten Wiesel]] in their work with kittens. The experiment involved sewing one eye shut and recording the cortical brain maps. Hubel and Wiesel saw that the portion of the kitten's brain associated with the shut eye was not idle, as expected. Instead, it processed visual information from the open eye. It was "…as though the brain didn't want to waste any 'cortical real estate' and had found a way to rewire itself." [44] => [45] => This implied neuroplasticity during the [[critical period]]. However, Merzenich argued that neuroplasticity could occur beyond the critical period. His first encounter with adult plasticity came when he was engaged in a postdoctoral study with Clinton Woosley. The experiment was based on observation of what occurred in the brain when one peripheral nerve was cut and subsequently regenerated. The two scientists micromapped the hand maps of monkey brains before and after cutting a peripheral nerve and sewing the ends together. Afterwards, the hand map in the brain that they expected to be jumbled was nearly normal. This was a substantial breakthrough. Merzenich asserted that, "If the brain map could normalize its structure in response to abnormal input, the prevailing view that we are born with a hardwired system had to be wrong. The brain had to be plastic." Merzenich received the 2016 [[Kavli Prize]] in Neuroscience "for the discovery of mechanisms that allow experience and neural activity to remodel brain function."{{Cite web| url=http://www.kavliprize.org/prizes-and-laureates/prizes/2016-kavli-prize-neuroscience| title=2016 Kavli Prize in Neuroscience| date=2016-06-02| access-date=2 June 2016| archive-date=5 June 2016| archive-url=https://web.archive.org/web/20160605124441/http://www.kavliprize.org/prizes-and-laureates/prizes/2016-kavli-prize-neuroscience| url-status=live}} [46] => [47] => == Neurobiology == [48] => [49] => There are different ideas and theories on what biological process allow for neuroplasticity to occur. The core of this phenomenon is based upon synapses and how connections between them change based on neuron functioning. It is widely agreed upon that neuroplasticity takes on many forms, as it is a result of a variety of pathways. These pathways, mainly signaling cascades, allow for gene expression alterations that lead to neuronal changes, and thus neuroplasticity. [50] => [51] => There are a number of other factors that are thought to play a role in the biological processes underlying the changing of neural networks in the brain. Some of these factors include synapse regulation via [[phosphorylation]], the role of inflammation and inflammatory cytokines, proteins such as Bcl-2 proteins and neutrophorins, and energy production via [[mitochondria]].{{Cite journal |last=Gulyaeva |first=N. V. |date=March 2017 |title=Molecular mechanisms of neuroplasticity: An expanding universe |url=http://link.springer.com/10.1134/S0006297917030014 |journal=Biochemistry (Moscow) |language=en |volume=82 |issue=3 |pages=237–242 |doi=10.1134/S0006297917030014 |pmid=28320264 |s2cid=6539117 |issn=0006-2979}} [52] => [53] => JT Wall and J Xu have traced the mechanisms underlying neuroplasticity. Re-organization is not cortically [[emergence|emergent]], but occurs at every level in the processing hierarchy; this produces the map changes observed in the cerebral cortex. [54] => [55] => == Types == [56] => [57] => Christopher Shaw and Jill McEachern (eds) in "Toward a theory of Neuroplasticity", state that there is no all-inclusive theory that overarches different frameworks and systems in the study of neuroplasticity. However, researchers often describe neuroplasticity as "the ability to make adaptive changes related to the structure and function of the nervous system."{{cite journal | vauthors = Zilles K | title = Neuronal plasticity as an adaptive property of the central nervous system | journal = Annals of Anatomy - Anatomischer Anzeiger | volume = 174 | issue = 5 | pages = 383–391 | date = October 1992 | pmid = 1333175 | doi = 10.1016/s0940-9602(11)80255-4 }} Correspondingly, two types of neuroplasticity are often discussed: structural neuroplasticity and functional neuroplasticity. [58] => [59] => ===Structural neuroplasticity=== [60] => [61] => Structural plasticity is often understood as the brain's ability to change its neuronal connections. New neurons are constantly produced and integrated into the central nervous system throughout the life span based on this type of neuroplasticity.{{cite book |last1=Puderbaugh |first1=Matt |last2=Emmady |first2=Prabhu D. |title=StatPearls |date=2023 |publisher=StatPearls Publishing |url=https://www.ncbi.nlm.nih.gov/books/NBK557811/ |access-date=10 October 2023 |chapter=Neuroplasticity|pmid=32491743 }} Researchers nowadays use multiple cross-sectional imaging methods (i.e. [[magnetic resonance imaging]] (MRI), [[computerized tomography]] (CT)) to study the structural alterations of the human brains.{{cite journal | vauthors = Chang Y | title = Reorganization and plastic changes of the human brain associated with skill learning and expertise | journal = Frontiers in Human Neuroscience | volume = 8 | issue = 55 | pages = 35 | date = 2014 | pmid = 24550812 | pmc = 3912552 | doi = 10.3389/fnhum.2014.00035 | doi-access = free }} This type of neuroplasticity often studies the effect of various internal or external stimuli on the brain's anatomical reorganization. The changes of [[grey matter]] proportion or the synaptic strength in the brain are considered as examples of structural neuroplasticity. Structural neuroplasticity is currently investigated more within the field of neuroscience in current academia. [62] => [63] => ===Functional neuroplasticity=== [64] => [65] => Functional plasticity refers to the brain's ability to alter and adapt the functional properties of neurons. Functional plasticity can occur in four known ways namely homologous area adaptation, map expansion, cross- model reassignment, and compensatory masquerade. Through homologous area adaptation a cognitive task is shifted from a damaged part of the brain to its homologous area in the brain. Functional changes like this occur usually in children rather than adults. In map expansion, cortical maps related to particular cognitive tasks expand due to frequent exposure to stimuli. Map expansion has been proven through experiments performed in relation to the study: experiment on effect of frequent stimulus on functional connectivity of the brain was observed in individuals learning spatial routes.{{Cite journal |last1=Keller |first1=Timothy A. |last2=Just |first2=Marcel Adam |date=2016-01-15 |title=Structural and functional neuroplasticity in human learning of spatial routes |journal=NeuroImage |language=en |volume=125 |pages=256–266 |doi=10.1016/j.neuroimage.2015.10.015 |pmid=26477660 |s2cid=2784354 |issn=1053-8119|doi-access=free }} Cross-model reassignment involves reception of novel input signals to a brain region which has been stripped off its default input. Functional plasticity through compensatory masquerade occurs using different cognitive processes for an already established cognitive task. [66] => [67] => The changes can occur in response to previous activity ([[activity-dependent plasticity]]) to acquire memory or in response to malfunction or damage of neurons ([[maladaptive plasticity]]) to compensate a pathological event. In the latter case the functions from one part of the brain transfer to another part of the brain based on the demand to produce recovery of behavioral or physiological processes.{{cite journal | vauthors = Freed WJ, de Medinaceli L, Wyatt RJ | title = Promoting functional plasticity in the damaged nervous system | journal = Science | volume = 227 | issue = 4694 | pages = 1544–1552 | date = March 1985 | pmid = 3975624 | doi = 10.1126/science.3975624 | bibcode = 1985Sci...227.1544F }} Regarding physiological forms of activity-dependent plasticity, those involving synapses are referred to as [[synaptic plasticity]]. The strengthening or weakening of synapses that results in an increase or decrease of firing rate of the neurons are called [[long-term potentiation]] (LTP) and [[long-term depression]] (LTD), respectively, and they are considered as examples of synaptic plasticity that are associated with memory.{{cite journal | vauthors = Patten AR, Yau SY, Fontaine CJ, Meconi A, Wortman RC, Christie BR | title = The Benefits of Exercise on Structural and Functional Plasticity in the Rodent Hippocampus of Different Disease Models | journal = Brain Plasticity | volume = 1 | issue = 1 | pages = 97–127 | date = October 2015 | pmid = 29765836 | pmc = 5928528 | doi = 10.3233/BPL-150016 }} The [[cerebellum]] is a typical structure with combinations of LTP/LTD and redundancy within the circuitry, allowing plasticity at several sites.{{cite journal | vauthors = Mitoma H, Kakei S, Yamaguchi K, Manto M | title = Physiology of Cerebellar Reserve: Redundancy and Plasticity of a Modular Machine | journal = International Journal of Molecular Sciences | volume = 22 | issue = 9 | pages = 4777 | date = April 2021 | pmid = 33946358 | pmc = 8124536 | doi = 10.3390/ijms22094777 | doi-access = free }} More recently it has become clearer that synaptic plasticity can be complemented by another form of activity-dependent plasticity involving the intrinsic excitability of neurons, which is referred to as [[intrinsic plasticity]].{{cite journal | vauthors = Zhang W, Linden DJ | title = The other side of the engram: experience-driven changes in neuronal intrinsic excitability | journal = Nature Reviews. Neuroscience | volume = 4 | issue = 11 | pages = 885–900 | date = November 2003 | pmid = 14595400 | doi = 10.1038/nrn1248 | s2cid = 17397545 }}{{cite journal | vauthors = Debanne D, Inglebert Y, Russier M | title = Plasticity of intrinsic neuronal excitability | journal = Current Opinion in Neurobiology | volume = 54 | pages = 73–82 | date = February 2019 | pmid = 30243042 | doi = 10.1016/j.conb.2018.09.001 | url = https://hal-amu.archives-ouvertes.fr/hal-01963474/file/Debannne-Russier-2019.pdf | access-date = 29 February 2020 | url-status = live | s2cid = 52812190 | archive-url = https://web.archive.org/web/20200203132111/https://hal-amu.archives-ouvertes.fr/hal-01963474/file/Debannne-Russier-2019.pdf | archive-date = 3 February 2020 }}{{cite journal | vauthors = Scheler, Gabriele | title = Learning intrinsic excitability in medium spiny neurons | journal = F1000Research | volume = 2 | pages = 88 | date = 2013 | pmid = 25520776 | doi = 10.12688/f1000research.2-88.v2| pmc = 4264637 | doi-access = free }} This, as opposed to [[homeostatic plasticity]] does not necessarily maintain the overall activity of a neuron within a network but contributes to encoding memories.{{cite journal | vauthors = Grasselli G, Boele HJ, Titley HK, Bradford N, van Beers L, Jay L, Beekhof GC, Busch SE, De Zeeuw CI, Schonewille M, Hansel C | title = SK2 channels in cerebellar Purkinje cells contribute to excitability modulation in motor-learning-specific memory traces | journal = PLOS Biology | volume = 18 | issue = 1 | pages = e3000596 | date = January 2020 | pmid = 31905212 | pmc = 6964916 | doi = 10.1371/journal.pbio.3000596 | doi-access = free }} Also, many studies have indicated functional neuroplasticity in the level of brain networks, where training alters the strength of functional connections.{{cite journal | vauthors = Duru AD, Balcioglu TH | title = Functional and Structural Plasticity of Brain in Elite Karate Athletes | journal = Journal of Healthcare Engineering | volume = 2018 | pages = 8310975 | date = 2018 | pmid = 30425820 | pmc = 6218732 | doi = 10.1155/2018/8310975 | doi-access = free }}{{cite journal | vauthors = Kelly C, Castellanos FX | title = Strengthening connections: functional connectivity and brain plasticity | journal = Neuropsychology Review | volume = 24 | issue = 1 | pages = 63–76 | date = March 2014 | pmid = 24496903 | pmc = 4059077 | doi = 10.1007/s11065-014-9252-y }} Although a recent study discusses that these observed changes should not directly relate to neuroplasticity, since they may root in the systematic requirement of the brain network for reorganization.{{cite journal | vauthors = Saberi M, Khosrowabadi R, Khatibi A, Misic B, Jafari G | title = Requirement to change of functional brain network across the lifespan | journal = PLOS ONE | volume = 16 | issue = 11 | pages = e0260091 | date = 2021 | pmid = 34793536 | pmc = 8601519 | doi = 10.1371/journal.pone.0260091 | bibcode = 2021PLoSO..1660091S | doi-access = free }} [68] => [69] => == Applications and examples == [70] => [71] => The adult brain is not entirely "hard-wired" with fixed [[neuronal circuit]]s. There are many instances of cortical and subcortical rewiring of neuronal circuits in response to training as well as in response to injury. [72] => [73] => There is ample evidence{{Cite journal |last1=Yu |first1=Feng |last2=Jiang |first2=Qing-jun |last3=Sun |first3=Xi-yan |last4=Zhang |first4=Rong-wei |date=2014-08-22 |title=A new case of complete primary cerebellar agenesis: clinical and imaging findings in a living patient |url=https://doi.org/10.1093/brain/awu239 |journal=Brain |volume=138 |issue=6 |pages=e353 |doi=10.1093/brain/awu239 |issn=0006-8950 |pmc=4614135 |pmid=25149410}} for the active, experience-dependent re-organization of the synaptic networks of the brain involving multiple inter-related structures including the cerebral cortex.{{Cite journal |last1=Scheler |first1=Gabriele |date=January 2023 |title=Sketch of a novel approach to a neural model [74] => |arxiv=2209.06865 |language=en }} The specific details of how this process occurs at the molecular and [[ultrastructure|ultrastructural]] levels are topics of active neuroscience research. The way experience can influence the synaptic organization of the brain is also the basis for a number of theories of brain function including the general theory of mind and [[neural Darwinism]]. The concept of neuroplasticity is also central to theories of memory and learning that are associated with experience-driven alteration of synaptic structure and function in studies of [[classical conditioning]] in invertebrate animal models such as ''[[Aplysia]]''. [75] => [76] => There is evidence that [[neurogenesis]] (birth of brain cells) occurs in the adult, rodent brain—and such changes can persist well into old age.{{Cite journal |last1=Duque |first1=Alvaro |last2=Arellano |first2=Jon I. |last3=Rakic |first3=Pasko |date=January 2022 |title=An assessment of the existence of adult neurogenesis in humans and value of its rodent models for neuropsychiatric diseases |journal=Molecular Psychiatry |language=en |volume=27 |issue=1 |pages=377–382 |doi=10.1038/s41380-021-01314-8 |pmid=34667259 |pmc=8967762 |issn=1476-5578}} The evidence for neurogenesis is mainly restricted to the [[hippocampus]] and [[olfactory bulb]], but research has revealed that other parts of the brain, including the cerebellum, may be involved as well. However, the degree of rewiring induced by the integration of new neurons in the established circuits is not known, and such rewiring may well be functionally redundant.{{cite journal |vauthors=França TF |date=November 2018 |title=Plasticity and redundancy in the integration of adult born neurons in the hippocampus |journal=Neurobiology of Learning and Memory |volume=155 |pages=136–142 |doi=10.1016/j.nlm.2018.07.007 |pmid=30031119 |doi-access=free |s2cid=51710989}} [77] => [78] => === Treatment of brain damage === [79] => [80] => A surprising consequence of neuroplasticity is that the brain activity associated with a given function can be transferred to a different location; this can result from normal experience and also occurs in the process of recovery from brain injury. Neuroplasticity is the fundamental issue that supports the scientific basis for treatment of [[acquired brain injury]] with goal-directed experiential therapeutic programs in the context of [[Rehabilitation (neuropsychology)|rehabilitation]] approaches to the functional consequences of the injury. [81] => [82] => Neuroplasticity is gaining popularity as a theory that, at least in part, explains improvements in functional outcomes with physical therapy post-stroke. Rehabilitation techniques that are supported by evidence which suggest cortical reorganization as the mechanism of change include [[constraint-induced movement therapy]], [[functional electrical stimulation]], treadmill training with body-weight support, and [[virtual reality therapy]]. [[Robotic rehabilitation|Robot assisted therapy]] is an emerging technique, which is also hypothesized to work by way of neuroplasticity, though there is currently insufficient evidence to determine the exact mechanisms of change when using this method.{{cite journal | vauthors = Young JA, Tolentino M | title = Neuroplasticity and its applications for rehabilitation | journal = American Journal of Therapeutics | volume = 18 | issue = 1 | pages = 70–80 | date = January 2011 | pmid = 21192249 | doi = 10.1097/MJT.0b013e3181e0f1a4 }} [83] => [84] => One group has developed a treatment that includes increased levels of [[progesterone]] injections in brain-injured patients. "Administration of progesterone after traumatic brain injury[https://web.archive.org/web/20060626124922/http://www.whsc.emory.edu/press_releases_video.cfm?id=brain_trauma Traumatic Brain Injury] (a story of TBI and the results of ProTECT using progesterone treatments) Emory University News Archives (TBI) and stroke reduces [[edema]], inflammation, and neuronal cell death, and enhances spatial reference memory and sensory-motor recovery."{{cite journal | vauthors = Cutler SM, Pettus EH, Hoffman SW, Stein DG | title = Tapered progesterone withdrawal enhances behavioral and molecular recovery after traumatic brain injury | journal = Experimental Neurology | volume = 195 | issue = 2 | pages = 423–429 | date = October 2005 | pmid = 16039652 | doi = 10.1016/j.expneurol.2005.06.003 | s2cid = 6305569 }} In a clinical trial, a group of severely injured patients had a 60% reduction in mortality after three days of progesterone injections. However, a study published in the ''[[The New England Journal of Medicine|New England Journal of Medicine]]'' in 2014 detailing the results of a multi-center NIH-funded phase III clinical trial of 882 patients found that treatment of acute traumatic brain injury with the hormone progesterone provides no significant benefit to patients when compared with placebo.{{cite web | url = http://news.emory.edu/stories/2014/12/protectiii_results_progesterone_tbi/ | title = Progesterone offers no significant benefit in traumatic brain injury clinical trial | archive-url = https://web.archive.org/web/20150327011928/http://news.emory.edu/stories/2014/12/protectiii_results_progesterone_tbi/ | archive-date = 27 March 2015 | publisher = Emory University | location = Atlanta, GA }} [85] => [86] => === Binocular vision === [87] => For decades, researchers assumed that humans had to acquire [[binocular vision]], in particular [[stereopsis]], in early childhood or they would never gain it. In recent years, however, successful improvements in persons with [[amblyopia]], [[convergence insufficiency]] or other stereo vision anomalies have become prime examples of neuroplasticity; binocular vision improvements and [[stereopsis recovery]] are now active areas of scientific and clinical research.{{cite journal | vauthors = Maino DM | title = Neuroplasticity: Teaching an old brain new tricks. | journal = Review of Optometry | date = January 2009 | volume = 39 | page = 46 | url = http://www.revoptom.com/continuing_education/tabviewtest/lessonid/106025/dnnprintmode/true/skinsrc/ | archive-url = https://web.archive.org/web/20140819102836/http://www.revoptom.com/continuing_education/tabviewtest/lessonid/106025/dnnprintmode/true/skinsrc/ | archive-date = 19 August 2014 }}{{cite journal | vauthors = Vedamurthy I, Huang SJ, Levi DM, Bavelier D, Knill DC |title=Recovery of stereopsis in adults through training in a virtual reality task|journal=Journal of Vision|date=27 December 2012|volume=12|number=14| page = 53 |doi=10.1167/12.14.53|doi-access=free}}{{cite journal | vauthors = Hess RF, Thompson B | title = New insights into amblyopia: binocular therapy and noninvasive brain stimulation | journal = Journal of AAPOS | volume = 17 | issue = 1 | pages = 89–93 | date = February 2013 | pmid = 23352385 | doi = 10.1016/j.jaapos.2012.10.018 }} [88] => [89] => ===Phantom limbs=== [90] => [[File:Mirror-box-comic.jpg|thumb|right|200px|A diagrammatic explanation of the mirror box. The patient places the intact limb into one side of the box (in this case the right hand) and the amputated limb into the other side. Due to the mirror, the patient sees a reflection of the intact hand where the missing limb would be (indicated in lower contrast). The patient thus receives artificial visual feedback that the "resurrected" limb is now moving when they move the good hand.]] [91] => {{Main|Phantom limb|Mirror box}} [92] => [93] => In the phenomenon of [[phantom limb]] sensation, a person continues to feel pain or sensation within a part of their body that has been [[Amputation|amputated]]. This is strangely common, occurring in 60–80% of amputees.{{cite journal | vauthors = Beaumont G, Mercier C, Michon PE, Malouin F, Jackson PL | title = Decreasing phantom limb pain through observation of action and imagery: a case series | journal = Pain Medicine | volume = 12 | issue = 2 | pages = 289–299 | date = February 2011 | pmid = 21276185 | doi = 10.1111/j.1526-4637.2010.01048.x | doi-access = free }} An [[Phantom limb#Neurological basis|explanation]] for this is based on the concept of neuroplasticity, as the [[cortical map]]s of the removed limbs are believed to have become engaged with the area around them in the [[postcentral gyrus]]. This results in activity within the surrounding area of the cortex being misinterpreted by the area of the cortex formerly responsible for the amputated limb. [94] => [95] => The relationship between phantom limb sensation and neuroplasticity is a complex one. In the early 1990s [[V.S. Ramachandran]] theorized that phantom limbs were the result of [[cortical remapping]]. However, in 1995 Herta Flor and her colleagues demonstrated that cortical remapping occurs only in patients who have phantom pain.{{cite journal | vauthors = Flor H, Elbert T, Knecht S, Wienbruch C, Pantev C, Birbaumer N, Larbig W, Taub E | title = Phantom-limb pain as a perceptual correlate of cortical reorganization following arm amputation | journal = Nature | volume = 375 | issue = 6531 | pages = 482–484 | date = June 1995 | pmid = 7777055 | doi = 10.1038/375482a0 | url = http://nbn-resolving.de/urn:nbn:de:bsz:352-opus-63710 | access-date = 21 December 2018 | url-status = live | s2cid = 205025856 | bibcode = 1995Natur.375..482F | archive-url = https://web.archive.org/web/20201120201812/http://kops.uni-konstanz.de/handle/123456789/11254 | archive-date = 20 November 2020 }} Her research showed that phantom limb pain (rather than referred sensations) was the perceptual correlate of cortical reorganization.{{cite journal | vauthors = Flor H | title = Cortical reorganisation and chronic pain: implications for rehabilitation | journal = Journal of Rehabilitation Medicine | volume = 35 | issue = 41 Suppl | pages = 66–72 | date = May 2003 | pmid = 12817660 | doi = 10.1080/16501960310010179 | doi-access = free }} This phenomenon is sometimes referred to as maladaptive plasticity. [96] => [97] => In 2009, Lorimer Moseley and Peter Brugger carried out an experiment in which they encouraged arm amputee subjects to use visual imagery to contort their phantom limbs into impossible{{Clarify|reason=Explain what is meant by 'impossible'|date=July 2020}} configurations. Four of the seven subjects succeeded in performing impossible movements of the phantom limb. This experiment suggests that the subjects had modified the neural representation of their phantom limbs and generated the motor commands needed to execute impossible movements in the absence of feedback from the body.{{cite journal | vauthors = Moseley GL, Brugger P | title = Interdependence of movement and anatomy persists when amputees learn a physiologically impossible movement of their phantom limb | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 106 | issue = 44 | pages = 18798–18802 | date = November 2009 | pmid = 19858475 | pmc = 2774040 | doi = 10.1073/pnas.0907151106 | doi-access = free | bibcode = 2009PNAS..10618798M }} The authors stated that: "In fact, this finding extends our understanding of the brain's plasticity because it is evidence that profound changes in the mental representation of the body can be induced purely by internal brain mechanisms—the brain truly does change itself." [98] => [99] => ===Chronic pain=== [100] => {{Main|Chronic pain}} [101] => Individuals who have chronic pain experience prolonged pain at sites that may have been previously injured, yet are otherwise currently healthy. This phenomenon is related to neuroplasticity due to a maladaptive reorganization of the nervous system, both peripherally and centrally. During the period of tissue damage, [[Noxious stimulus|noxious stimuli]] and [[inflammation]] cause an elevation of nociceptive input from the periphery to the central nervous system. Prolonged [[nociception]] from the periphery then elicits a neuroplastic response at the cortical level to change its [[Somatotopic arrangement|somatotopic organization]] for the painful site, inducing [[central sensitization]].{{cite journal | vauthors = Seifert F, Maihöfner C | title = Functional and structural imaging of pain-induced neuroplasticity | journal = Current Opinion in Anesthesiology | volume = 24 | issue = 5 | pages = 515–523 | date = October 2011 | pmid = 21822136 | doi = 10.1097/aco.0b013e32834a1079 | s2cid = 6680116 }} For instance, individuals experiencing [[complex regional pain syndrome]] demonstrate a diminished cortical somatotopic representation of the hand contralaterally as well as a decreased spacing between the hand and the mouth.{{cite journal | vauthors = Maihöfner C, Handwerker HO, Neundörfer B, Birklein F | title = Patterns of cortical reorganization in complex regional pain syndrome | journal = Neurology | volume = 61 | issue = 12 | pages = 1707–1715 | date = December 2003 | pmid = 14694034 | doi = 10.1212/01.wnl.0000098939.02752.8e | s2cid = 23080189 }} Additionally, chronic pain has been reported to significantly reduce the volume of [[grey matter]] in the brain globally, and more specifically at the [[prefrontal cortex]] and right [[thalamus]].{{cite journal | vauthors = Apkarian AV, Sosa Y, Sonty S, Levy RM, Harden RN, Parrish TB, Gitelman DR | title = Chronic back pain is associated with decreased prefrontal and thalamic gray matter density | journal = The Journal of Neuroscience | volume = 24 | issue = 46 | pages = 10410–10415 | date = November 2004 | pmid = 15548656 | pmc = 6730296 | doi = 10.1523/JNEUROSCI.2541-04.2004 | url = https://zenodo.org/record/1008676 | access-date = 8 September 2019 | url-status = live | archive-url = https://web.archive.org/web/20200622091051/https://zenodo.org/record/1008676 | archive-date = 22 June 2020 }} However, following treatment, these abnormalities in cortical reorganization and grey matter volume are resolved, as well as their symptoms. Similar results have been reported for phantom limb pain,{{cite journal | vauthors = Karl A, Birbaumer N, Lutzenberger W, Cohen LG, Flor H | title = Reorganization of motor and somatosensory cortex in upper extremity amputees with phantom limb pain | journal = The Journal of Neuroscience | volume = 21 | issue = 10 | pages = 3609–3618 | date = May 2001 | pmid = 11331390 | pmc = 6762494 | doi = 10.1523/JNEUROSCI.21-10-03609.2001 }} [[Low back pain#Chronic pain|chronic low back pain]]{{cite journal | vauthors = Flor H, Braun C, Elbert T, Birbaumer N | title = Extensive reorganization of primary somatosensory cortex in chronic back pain patients | journal = Neuroscience Letters | volume = 224 | issue = 1 | pages = 5–8 | date = March 1997 | pmid = 9132689 | doi = 10.1016/s0304-3940(97)13441-3 | url = http://nbn-resolving.de/urn:nbn:de:bsz:352-opus-41386 | access-date = 21 December 2018 | url-status = live | s2cid = 18151663 | archive-url = https://web.archive.org/web/20201120201813/http://kops.uni-konstanz.de/handle/123456789/11263 | archive-date = 20 November 2020 }} and [[carpal tunnel syndrome]].{{cite journal | vauthors = Napadow V, Kettner N, Ryan A, Kwong KK, Audette J, Hui KK | title = Somatosensory cortical plasticity in carpal tunnel syndrome--a cross-sectional fMRI evaluation | journal = NeuroImage | volume = 31 | issue = 2 | pages = 520–530 | date = June 2006 | pmid = 16460960 | doi = 10.1016/j.neuroimage.2005.12.017 | s2cid = 7367285 }} [102] => [103] => ===Meditation=== [104] => {{Main|Research on meditation}} [105] => A number of studies have linked meditation practice to differences in cortical thickness or density of [[gray matter]].{{cite journal | vauthors = Sasmita AO, Kuruvilla J, Ling AP | title = Harnessing neuroplasticity: modern approaches and clinical future | journal = The International Journal of Neuroscience | volume = 128 | issue = 11 | pages = 1061–1077 | date = November 2018 | pmid = 29667473 | doi = 10.1080/00207454.2018.1466781 | s2cid = 4957270 }}{{cite journal | vauthors = Pagnoni G, Cekic M | title = Age effects on gray matter volume and attentional performance in Zen meditation | journal = Neurobiology of Aging | volume = 28 | issue = 10 | pages = 1623–1627 | date = October 2007 | pmid = 17655980 | doi = 10.1016/j.neurobiolaging.2007.06.008 | s2cid = 16755503 | hdl = 11380/609140 }}{{cite journal | vauthors = Vestergaard-Poulsen P, van Beek M, Skewes J, Bjarkam CR, Stubberup M, Bertelsen J, Roepstorff A | title = Long-term meditation is associated with increased gray matter density in the brain stem | journal = NeuroReport | volume = 20 | issue = 2 | pages = 170–174 | date = January 2009 | pmid = 19104459 | doi = 10.1097/WNR.0b013e328320012a | s2cid = 14263267 }}{{cite journal | vauthors = Luders E, Toga AW, Lepore N, Gaser C | title = The underlying anatomical correlates of long-term meditation: larger hippocampal and frontal volumes of gray matter | journal = NeuroImage | volume = 45 | issue = 3 | pages = 672–678 | date = April 2009 | pmid = 19280691 | pmc = 3184843 | doi = 10.1016/j.neuroimage.2008.12.061 }} One of the most well-known studies to demonstrate this was led by [[Sara Lazar]], from Harvard University, in 2000.{{cite journal | vauthors = Lazar SW, Kerr CE, Wasserman RH, Gray JR, Greve DN, Treadway MT, McGarvey M, Quinn BT, Dusek JA, Benson H, Rauch SL, Moore CI, Fischl B | title = Meditation experience is associated with increased cortical thickness | journal = NeuroReport | volume = 16 | issue = 17 | pages = 1893–1897 | date = November 2005 | pmid = 16272874 | pmc = 1361002 | doi = 10.1097/01.wnr.0000186598.66243.19 }} [[Richard Davidson]], a neuroscientist at the [[University of Wisconsin]], has led experiments in collaboration with the [[Dalai Lama]] on effects of meditation on the brain. His results suggest that meditation may lead to change in the physical structure of brain regions associated with [[attention]], [[anxiety]], [[Depression (mood)|depression]], [[fear]], [[anger]], and compassion as well as the ability of the body to heal itself.{{cite journal | vauthors = Lutz A, Greischar LL, Rawlings NB, Ricard M, Davidson RJ | title = Long-term meditators self-induce high-amplitude gamma synchrony during mental practice | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 101 | issue = 46 | pages = 16369–16373 | date = November 2004 | pmid = 15534199 | pmc = 526201 | doi = 10.1073/pnas.0407401101 | doi-access = free | bibcode = 2004PNAS..10116369L }}{{cite journal | vauthors = Davidson RJ, Lutz A | title = Buddha's Brain: Neuroplasticity and Meditation | journal = IEEE Signal Processing Magazine | volume = 25 | issue = 1 | pages = 176–174 | date = January 2008 | pmid = 20871742 | pmc = 2944261 | doi = 10.1109/MSP.2008.4431873 | url = https://centerhealthyminds.org/assets/files-publications/DavidsonBuddhaIEEESignalProcessingMagazine.pdf | access-date = 19 April 2018 | url-status = live | bibcode = 2008ISPM...25..176D | archive-url = https://web.archive.org/web/20120112084117/http://brainimaging.waisman.wisc.edu/publications/2008/DavidsonBuddhaIEEE.pdf | archive-date = 12 January 2012 }} [106] => [107] => === Artistic engagement and art therapy === [108] => There is substantial evidence that artistic engagement in a therapeutic environment can create changes in neural network connections as well as increase cognitive flexibility.{{Cite journal |last1=Lin |first1=Chia-Shu |last2=Liu |first2=Yong |last3=Huang |first3=Wei-Yuan |last4=Lu |first4=Chia-Feng |last5=Teng |first5=Shin |last6=Ju |first6=Tzong-Ching |last7=He |first7=Yong |last8=Wu |first8=Yu-Te |last9=Jiang |first9=Tianzi |last10=Hsieh |first10=Jen-Chuen |date=2013 |title=Sculpting the Intrinsic Modular Organization of Spontaneous Brain Activity by Art |journal=PLOS ONE |volume=8 |issue=6 |pages=e66761 |doi=10.1371/journal.pone.0066761 |issn=1932-6203 |pmc=3694132 |pmid=23840527|bibcode=2013PLoSO...866761L |doi-access=free }}{{Cite journal |last=Patel |first=Aniruddh D. |date=July 2003 |title=Language, music, syntax and the brain |url=https://www.nature.com/articles/nn1082 |journal=Nature Neuroscience |language=en |volume=6 |issue=7 |pages=674–681 |doi=10.1038/nn1082 |pmid=12830158 |s2cid=15689983 |issn=1546-1726}} In one 2013 study, researchers found evidence that long-term, habitual artistic training (e.g. musical instrument practice, purposeful painting, etc.) can "macroscopically imprint a neural network system of spontaneous activity in which the related brain regions become functionally and topologically modularized in both domain-general and domain-specific manners".{{Cite journal |last1=Lin |first1=Chia-Shu |last2=Liu |first2=Yong |last3=Huang |first3=Wei-Yuan |last4=Lu |first4=Chia-Feng |last5=Teng |first5=Shin |last6=Ju |first6=Tzong-Ching |last7=He |first7=Yong |last8=Wu |first8=Yu-Te |last9=Jiang |first9=Tianzi |last10=Hsieh |first10=Jen-Chuen |date=2013-06-26 |title=Sculpting the Intrinsic Modular Organization of Spontaneous Brain Activity by Art |journal=PLOS ONE |language=en |volume=8 |issue=6 |pages=e66761 |doi=10.1371/journal.pone.0066761 |issn=1932-6203 |pmc=3694132 |pmid=23840527|bibcode=2013PLoSO...866761L |doi-access=free }} In simple terms, brains repeatedly exposed to artistic training over long periods develop adaptations to make such activity both easier and more likely to spontaneously occur. [109] => [110] => Some researchers and academics have suggested that artistic engagement has substantially altered the human brain throughout our evolutionary history. D.W Zaidel, adjunct professor of behavioral neuroscience and contributor at [[VAGA]], has written that "evolutionary theory links the symbolic nature of art to critical pivotal brain changes in ''Homo sapiens'' supporting increased development of language and hierarchical social grouping".{{Cite journal |last=Zaidel |first=Dahlia W |date=February 2010 |title=Art and brain: insights from neuropsychology, biology and evolution |journal=Journal of Anatomy |volume=216 |issue=2 |pages=177–183 |doi=10.1111/j.1469-7580.2009.01099.x |issn=0021-8782 |pmc=2815940 |pmid=19490399}} [111] => [112] => ===Fitness and exercise=== [113] => {{See also|Neurobiological effects of physical exercise#Structural growth}} [114] => Aerobic exercise increases the production of [[neurotrophic factors]] (compounds that promote growth or survival of neurons), such as [[brain-derived neurotrophic factor]] (BDNF), [[insulin-like growth factor 1]] (IGF-1), and [[vascular endothelial growth factor]] (VEGF).{{cite journal | vauthors = Tarumi T, Zhang R | title = Cerebral hemodynamics of the aging brain: risk of Alzheimer disease and benefit of aerobic exercise | journal = Frontiers in Physiology | volume = 5 | pages = 6 | date = January 2014 | pmid = 24478719 | pmc = 3896879 | doi = 10.3389/fphys.2014.00006 | quote = Exercise-related improvements in brain function and structure may be conferred by the concurrent adaptations in vascular function and structure. Aerobic exercise increases the peripheral levels of growth factors (e.g., BDNF, IFG-1, and VEGF) that cross the blood-brain barrier (BBB) and stimulate neurogenesis and angiogenesis (Trejo et al., 2001; Lee et al., 2002; Fabel et al., 2003; Lopez-Lopez et al., 2004). | doi-access = free }}{{cite journal | vauthors = Szuhany KL, Bugatti M, Otto MW | title = A meta-analytic review of the effects of exercise on brain-derived neurotrophic factor | journal = Journal of Psychiatric Research | volume = 60 | pages = 56–64 | date = January 2015 | pmid = 25455510 | pmc = 4314337 | doi = 10.1016/j.jpsychires.2014.10.003 | quote = Consistent evidence indicates that exercise improves cognition and mood, with preliminary evidence suggesting that brain-derived neurotrophic factor (BDNF) may mediate these effects. The aim of the current meta-analysis was to provide an estimate of the strength of the association between exercise and increased BDNF levels in humans across multiple exercise paradigms. We conducted a meta-analysis of 29 studies (N = 1111 participants) examining the effect of exercise on BDNF levels in three exercise paradigms: (1) a single session of exercise, (2) a session of exercise following a program of regular exercise, and (3) resting BDNF levels following a program of regular exercise. Moderators of this effect were also examined. Results demonstrated a moderate effect size for increases in BDNF following a single session of exercise (Hedges' g = 0.46, p < 0.001). Further, regular exercise intensified the effect of a session of exercise on BDNF levels (Hedges' g = 0.59, p = 0.02). Finally, results indicated a small effect of regular exercise on resting BDNF levels (Hedges' g = 0.27, p = 0.005). ... Effect size analysis supports the role of exercise as a strategy for enhancing BDNF activity in humans }} Exercise-induced effects on the hippocampus are associated with measurable improvements in [[spatial memory]].{{cite journal | vauthors = Lees C, Hopkins J | title = Effect of aerobic exercise on cognition, academic achievement, and psychosocial function in children: a systematic review of randomized control trials | journal = Preventing Chronic Disease | volume = 10 | pages = E174 | date = October 2013 | pmid = 24157077 | pmc = 3809922 | doi = 10.5888/pcd10.130010 }}{{cite journal | vauthors = Carvalho A, Rea IM, Parimon T, Cusack BJ | title = Physical activity and cognitive function in individuals over 60 years of age: a systematic review | journal = Clinical Interventions in Aging | volume = 9 | pages = 661–682 | year = 2014 | pmid = 24748784 | pmc = 3990369 | doi = 10.2147/CIA.S55520 | doi-access = free }} Consistent aerobic exercise over a period of several months induces marked [[clinically significant]] improvements in [[executive function]] (i.e., the "[[cognitive control]]" of behavior) and increased [[gray matter]] volume in multiple brain regions, particularly those that give rise to cognitive control.{{cite journal | vauthors = Guiney H, Machado L | title = Benefits of regular aerobic exercise for executive functioning in healthy populations | journal = Psychonomic Bulletin & Review | volume = 20 | issue = 1 | pages = 73–86 | date = February 2013 | pmid = 23229442 | doi = 10.3758/s13423-012-0345-4 | doi-access = free }}{{cite journal | vauthors = Buckley J, Cohen JD, Kramer AF, McAuley E, Mullen SP | title = Cognitive control in the self-regulation of physical activity and sedentary behavior | journal = Frontiers in Human Neuroscience | volume = 8 | pages = 747 | year = 2014 | pmid = 25324754 | pmc = 4179677 | doi = 10.3389/fnhum.2014.00747 | doi-access = free }} The brain structures that show the greatest improvements in gray matter volume in response to aerobic exercise are the [[prefrontal cortex]] and [[hippocampus]]; moderate improvements are seen in the [[anterior cingulate cortex]], [[parietal cortex]], [[cerebellum]], [[caudate nucleus]], and [[nucleus accumbens]]. Higher [[physical fitness]] scores (measured by [[VO2 max|VO2 max]]) are associated with better executive function, faster processing speed, and greater volume of the hippocampus, caudate nucleus, and nucleus accumbens. [115] => [116] => === Deafness and loss of hearing === [117] => Due to hearing loss, the [[auditory cortex]] and other association areas of the brain in deaf and/or hard of hearing people undergo compensatory plasticity.{{cite journal | vauthors = Karns CM, Dow MW, Neville HJ | title = Altered cross-modal processing in the primary auditory cortex of congenitally deaf adults: a visual-somatosensory fMRI study with a double-flash illusion | journal = The Journal of Neuroscience | volume = 32 | issue = 28 | pages = 9626–9638 | date = July 2012 | pmid = 22787048 | pmc = 3752073 | doi = 10.1523/JNEUROSCI.6488-11.2012 | url = https://www.jneurosci.org/content/32/28/9626 | url-status = live | archive-url = https://web.archive.org/web/20200317151339/https://www.jneurosci.org/content/32/28/9626 | archive-date = 17 March 2020 }}{{cite journal | vauthors = Bottari D, Heimler B, Caclin A, Dalmolin A, Giard MH, Pavani F | title = Visual change detection recruits auditory cortices in early deafness | journal = NeuroImage | volume = 94 | pages = 172–184 | date = July 2014 | pmid = 24636881 | doi = 10.1016/j.neuroimage.2014.02.031 | url = http://www.sciencedirect.com/science/article/pii/S1053811914001463 | access-date = 11 November 2020 | url-status = live | s2cid = 207189746 | archive-url = https://web.archive.org/web/20201120201811/https://www.sciencedirect.com/science/article/abs/pii/S1053811914001463 | archive-date = 20 November 2020 }}{{cite journal | vauthors = Bavelier D, Brozinsky C, Tomann A, Mitchell T, Neville H, Liu G | title = Impact of early deafness and early exposure to sign language on the cerebral organization for motion processing | journal = The Journal of Neuroscience | volume = 21 | issue = 22 | pages = 8931–8942 | date = November 2001 | pmid = 11698604 | pmc = 6762265 | doi = 10.1523/JNEUROSCI.21-22-08931.2001 | url = https://www.jneurosci.org/content/21/22/8931 | url-status = live | archive-url = https://web.archive.org/web/20200604085354/https://www.jneurosci.org/content/21/22/8931 | archive-date = 4 June 2020 }} The auditory cortex is usually reserved for processing auditory information in hearing people now is redirected to serve other functions, especially for [[Visual system|vision]] and [[Somatosensory system|somatosensation]]. [118] => [119] => Deaf individuals have enhanced peripheral visual attention,{{cite journal | vauthors = Neville HJ, Lawson D | title = Attention to central and peripheral visual space in a movement detection task: an event-related potential and behavioral study. II. Congenitally deaf adults | journal = Brain Research | volume = 405 | issue = 2 | pages = 268–283 | date = March 1987 | pmid = 3567605 | doi = 10.1016/0006-8993(87)90296-4 | s2cid = 41719446 }} better motion change but not color change detection ability in visual tasks,{{cite journal | vauthors = Armstrong BA, Neville HJ, Hillyard SA, Mitchell TV | title = Auditory deprivation affects processing of motion, but not color | journal = Brain Research. Cognitive Brain Research | volume = 14 | issue = 3 | pages = 422–434 | date = November 2002 | pmid = 12421665 | doi = 10.1016/S0926-6410(02)00211-2 }} more effective visual search,{{cite journal | vauthors = Stivalet P, Moreno Y, Richard J, Barraud PA, Raphel C | title = Differences in visual search tasks between congenitally deaf and normally hearing adults | journal = Brain Research. Cognitive Brain Research | volume = 6 | issue = 3 | pages = 227–232 | date = January 1998 | pmid = 9479074 | doi = 10.1016/S0926-6410(97)00026-8 }} and faster response time for visual targets{{cite journal | vauthors = Heimler B, Pavani F | title = Response speed advantage for vision does not extend to touch in early deaf adults | journal = Experimental Brain Research | volume = 232 | issue = 4 | pages = 1335–1341 | date = April 2014 | pmid = 24477765 | doi = 10.1007/s00221-014-3852-x | url = http://link.springer.com/10.1007/s00221-014-3852-x | access-date = 11 November 2020 | url-status = live | hdl-access = free | s2cid = 18995518 | archive-date = 4 June 2018 | archive-url = https://web.archive.org/web/20180604000503/https://link.springer.com/article/10.1007%2Fs00221-014-3852-x | hdl = 11572/67241 }}{{cite journal | vauthors = Hauthal N, Debener S, Rach S, Sandmann P, Thorne JD | title = Visuo-tactile interactions in the congenitally deaf: a behavioral and event-related potential study | journal = Frontiers in Integrative Neuroscience | volume = 8 | pages = 98 | date = 2015 | pmid = 25653602 | pmc = 4300915 | doi = 10.3389/fnint.2014.00098 | doi-access = free }} compared to hearing individuals. Altered visual processing in deaf people is often found to be associated with the repurposing of other brain areas including [[Auditory cortex|primary auditory cortex]], [[Posterior parietal cortex|posterior parietal association cortex]] (PPAC), and [[anterior cingulate cortex]] (ACC).{{cite journal | vauthors = Scott GD, Karns CM, Dow MW, Stevens C, Neville HJ | title = Enhanced peripheral visual processing in congenitally deaf humans is supported by multiple brain regions, including primary auditory cortex | journal = Frontiers in Human Neuroscience | volume = 8 | pages = 177 | date = 2014 | pmid = 24723877 | pmc = 3972453 | doi = 10.3389/fnhum.2014.00177 | doi-access = free }} A review by Bavelier et al. (2006) summarizes many aspects on the topic of visual ability comparison between deaf and hearing individuals.{{cite journal | vauthors = Bavelier D, Dye MW, Hauser PC | title = Do deaf individuals see better? | journal = Trends in Cognitive Sciences | volume = 10 | issue = 11 | pages = 512–518 | date = November 2006 | pmid = 17015029 | pmc = 2885708 | doi = 10.1016/j.tics.2006.09.006 }} [120] => [121] => Brain areas that serve a function in auditory processing repurpose to process somatosensory information in congenitally deaf people. They have higher sensitivity in detecting frequency change in vibration above threshold{{cite journal | vauthors = Levänen S, Hamdorf D | title = Feeling vibrations: enhanced tactile sensitivity in congenitally deaf humans | journal = Neuroscience Letters | volume = 301 | issue = 1 | pages = 75–77 | date = March 2001 | pmid = 11239720 | doi = 10.1016/S0304-3940(01)01597-X | url = http://www.sciencedirect.com/science/article/pii/S030439400101597X | access-date = 11 November 2020 | url-status = live | s2cid = 1650771 | archive-url = https://web.archive.org/web/20201120201844/https://www.sciencedirect.com/science/article/abs/pii/S030439400101597X | archive-date = 20 November 2020 }} and higher and more widespread activation in auditory cortex under somatosensory stimulation.{{cite journal | vauthors = Auer ET, Bernstein LE, Sungkarat W, Singh M | title = Vibrotactile activation of the auditory cortices in deaf versus hearing adults | journal = NeuroReport | volume = 18 | issue = 7 | pages = 645–648 | date = May 2007 | pmid = 17426591 | pmc = 1934619 | doi = 10.1097/WNR.0b013e3280d943b9 | url = https://journals.lww.com/neuroreport/Abstract/2007/05070/Vibrotactile_activation_of_the_auditory_cortices.5.aspx | url-status = live | archive-url = https://web.archive.org/web/20201120201843/https://journals.lww.com/neuroreport/Abstract/2007/05070/Vibrotactile_activation_of_the_auditory_cortices.5.aspx | archive-date = 20 November 2020 }} However, speeded response for somatosensory stimuli is not found in deaf adults. [122] => [123] => ==== Cochlear implant ==== [124] => Neuroplasticity is involved in the development of sensory function. The brain is born immature and then adapts to sensory inputs after birth. In the auditory system, congenital hearing loss, a rather frequent inborn condition affecting 1 of 1000 newborns, has been shown to affect auditory development, and implantation of a [[cochlear implant|sensory prostheses]] activating the auditory system has prevented the deficits and induced functional maturation of the auditory system.{{cite journal | vauthors = Kral A, Sharma A | title = Developmental neuroplasticity after cochlear implantation | journal = Trends in Neurosciences | volume = 35 | issue = 2 | pages = 111–122 | date = February 2012 | pmid = 22104561 | pmc = 3561718 | doi = 10.1016/j.tins.2011.09.004 }} Due to a sensitive period for plasticity, there is also a sensitive period for such intervention within the first 2–4 years of life. Consequently, in prelingually deaf children, early [[cochlear implant]]ation, as a rule, allows the children to learn the mother language and acquire acoustic communication.{{cite journal | vauthors = Kral A, O'Donoghue GM | title = Profound deafness in childhood | journal = The New England Journal of Medicine | volume = 363 | issue = 15 | pages = 1438–1450 | date = October 2010 | pmid = 20925546 | doi = 10.1056/nejmra0911225 | s2cid = 13639137 }} [125] => [126] => === Blindness === [127] => Due to vision loss, the [[visual cortex]] in blind people may undergo cross-modal plasticity, and therefore other senses may have enhanced abilities. Or the opposite could occur, with the lack of visual input weakening the development of other sensory systems. One study suggests that the right posterior middle temporal gyrus and [[superior occipital gyrus]] reveal more activation in the blind than in the sighted people during a sound-moving detection task.{{cite journal | vauthors = Dormal G, Rezk M, Yakobov E, Lepore F, Collignon O | title = Auditory motion in the sighted and blind: Early visual deprivation triggers a large-scale imbalance between auditory and "visual" brain regions | journal = NeuroImage | volume = 134 | pages = 630–644 | date = July 2016 | pmid = 27107468 | doi = 10.1016/j.neuroimage.2016.04.027 | url = http://www.sciencedirect.com/science/article/pii/S1053811916300611 | access-date = 11 November 2020 | url-status = live | s2cid = 25832602 | archive-url = https://web.archive.org/web/20201120201844/https://www.sciencedirect.com/science/article/abs/pii/S1053811916300611 | archive-date = 20 November 2020 }} Several studies support the latter idea and found weakened ability in audio distance evaluation, proprioceptive reproduction, threshold for visual bisection, and judging minimum audible angle.{{cite journal | vauthors = Cappagli G, Cocchi E, Gori M | title = Auditory and proprioceptive spatial impairments in blind children and adults | journal = Developmental Science | volume = 20 | issue = 3 | pages = e12374 | date = May 2017 | pmid = 26613827 | doi = 10.1111/desc.12374 | url = http://doi.wiley.com/10.1111/desc.12374 | access-date = 11 November 2020 | url-status = live | archive-url = https://web.archive.org/web/20201120201854/https://onlinelibrary.wiley.com/doi/abs/10.1111/desc.12374 | archive-date = 20 November 2020 }}{{cite journal | vauthors = Vercillo T, Burr D, Gori M | title = Early visual deprivation severely compromises the auditory sense of space in congenitally blind children | journal = Developmental Psychology | volume = 52 | issue = 6 | pages = 847–853 | date = June 2016 | pmid = 27228448 | pmc = 5053362 | doi = 10.1037/dev0000103 }} [128] => [129] => ====Human echolocation==== [130] => [131] => [[Human echolocation]] is a learned ability for humans to sense their environment from echoes. This ability is used by some [[blindness|blind]] people to navigate their environment and sense their surroundings in detail. Studies in 2010{{cite journal | vauthors = Thaler L, Arnott SR, Goodale MA |title=Human Echolocation I |journal=Journal of Vision |date=13 August 2010 |volume=10 |issue=7 |pages=1050 |doi=10.1167/10.7.1050 |doi-access=free}} and 2011 using [[functional magnetic resonance imaging]] techniques have shown that parts of the brain associated with visual processing are adapted for the new skill of echolocation. Studies with blind patients, for example, suggest that the click-echoes heard by these patients were processed by brain regions devoted to vision rather than audition. [132] => [133] => === Attention deficit hyperactivity disorder === [134] => {{See also|Attention deficit hyperactivity disorder#Brain structure}} [135] => Reviews of MRI and electroencephalography (EEG) studies on individuals with ADHD suggest that the long-term treatment of ADHD with stimulants, such as [[amphetamine]] or [[methylphenidate]], decreases abnormalities in brain structure and function found in subjects with ADHD, and improves function in several parts of the brain, such as the right [[caudate nucleus]] of the [[basal ganglia]],{{cite journal | vauthors = Hart H, Radua J, Nakao T, Mataix-Cols D, Rubia K | title = Meta-analysis of functional magnetic resonance imaging studies of inhibition and attention in attention-deficit/hyperactivity disorder: exploring task-specific, stimulant medication, and age effects | journal = JAMA Psychiatry | volume = 70 | issue = 2 | pages = 185–198 | date = February 2013 | pmid = 23247506 | doi = 10.1001/jamapsychiatry.2013.277 | doi-access = }}{{cite journal | vauthors = Spencer TJ, Brown A, Seidman LJ, Valera EM, Makris N, Lomedico A, Faraone SV, Biederman J | title = Effect of psychostimulants on brain structure and function in ADHD: a qualitative literature review of magnetic resonance imaging-based neuroimaging studies | journal = The Journal of Clinical Psychiatry | volume = 74 | issue = 9 | pages = 902–917 | date = September 2013 | pmid = 24107764 | pmc = 3801446 | doi = 10.4088/JCP.12r08287 }}{{cite journal | vauthors = Frodl T, Skokauskas N | title = Meta-analysis of structural MRI studies in children and adults with attention deficit hyperactivity disorder indicates treatment effects | journal = Acta Psychiatrica Scandinavica | volume = 125 | issue = 2 | pages = 114–126 | date = February 2012 | pmid = 22118249 | doi = 10.1111/j.1600-0447.2011.01786.x | quote = Basal ganglia regions like the right globus pallidus, the right putamen, and the nucleus caudatus are structurally affected in children with ADHD. These changes and alterations in limbic regions like ACC and amygdala are more pronounced in non-treated populations and seem to diminish over time from child to adulthood. Treatment seems to have positive effects on brain structure. | s2cid = 25954331 | doi-access = free }} left [[ventrolateral prefrontal cortex]] (VLPFC), and [[superior temporal gyrus]].{{cite journal | vauthors = Kowalczyk OS, Cubillo AI, Smith A, Barrett N, Giampietro V, Brammer M, Simmons A, Rubia K | title = Methylphenidate and atomoxetine normalise fronto-parietal underactivation during sustained attention in ADHD adolescents | journal = European Neuropsychopharmacology | volume = 29 | issue = 10 | pages = 1102–1116 | date = October 2019 | pmid = 31358436 | doi = 10.1016/j.euroneuro.2019.07.139 | url = http://www.sciencedirect.com/science/article/pii/S0924977X19304365 | access-date = 11 November 2020 | url-status = live | s2cid = 198983414 | archive-url = https://web.archive.org/web/20201120201847/https://www.sciencedirect.com/science/article/abs/pii/S0924977X19304365 | archive-date = 20 November 2020 }} [136] => [137] => === In early child development === [138] => Neuroplasticity is most active in childhood as a part of normal [[human development (biology)|human development]], and can also be seen as an especially important mechanism for children in terms of risk and resiliency.{{cite journal | vauthors = Masten AS | title = Resilience in children threatened by extreme adversity: frameworks for research, practice, and translational synergy | journal = Development and Psychopathology | volume = 23 | issue = 2 | pages = 493–506 | date = May 2011 | pmid = 23786691 | doi = 10.1017/S0954579411000198 | s2cid = 12068256 }} Trauma is considered a great risk as it negatively affects many areas of the brain and puts a strain on the sympathetic nervous system from constant activation. Trauma thus alters the brain's connections such that children who have experienced trauma may be hyper vigilant or overly aroused.{{cite journal| vauthors = Schore AN |s2cid=9711339 |title=The effects of early relational trauma on right brain development, affect regulation, and infant mental health|journal=Infant Mental Health Journal|date=2001|volume=1|issue=2|pages=201–269|doi=10.1002/1097-0355(200101/04)22:1<201::AID-IMHJ8>3.0.CO;2-9}} However, a child's brain can cope with these adverse effects through the actions of neuroplasticity.{{cite book | vauthors = Cioni G, D'Acunto G, Guzzetta A | title = Gene Expression to Neurobiology and Behavior: Human Brain Development and Developmental Disorders | chapter = Perinatal brain damage in children | volume = 189 | pages = 139–154 | year = 2011 | pmid = 21489387 | doi = 10.1016/B978-0-444-53884-0.00022-1 | isbn = 978-0-444-53884-0 | series = Progress in Brain Research }} [139] => [140] => Neuroplasticity is shown in four different categories in children and covering a wide variety of neuronal functioning. These four types include impaired, excessive, adaptive, and plasticity.{{cite journal | vauthors = Mundkur N | title = Neuroplasticity in children | journal = Indian Journal of Pediatrics | volume = 72 | issue = 10 | pages = 855–857 | date = October 2005 | pmid = 16272658 | doi = 10.1007/BF02731115 | s2cid = 32108524 }} [141] => [142] => There are many examples of neuroplasticity in human development. For example, Justine Ker and Stephen Nelson looked at the effects of musical training on neuroplasticity, and found that musical training can contribute to experience dependent structural plasticity. This is when changes in the brain occur based on experiences that are unique to an individual. Examples of this are learning multiple languages, playing a sport, doing theatre, etc. A study done by Hyde in 2009, showed that changes in the brain of children could be seen in as little as 15 months of musical training.{{cite journal | vauthors = Hyde KL, Lerch J, Norton A, Forgeard M, Winner E, Evans AC, Schlaug G | title = Musical training shapes structural brain development | journal = The Journal of Neuroscience | volume = 29 | issue = 10 | pages = 3019–3025 | date = March 2009 | pmid = 19279238 | pmc = 2996392 | doi = 10.1523/JNEUROSCI.5118-08.2009 }} Ker and Nelson suggest this degree of plasticity in the brains of children can "help provide a form of intervention for children... with developmental disorders and neurological diseases."{{cite journal | vauthors = Ker J, Nelson S | title = The effects of musical training on brain plasticity and cognitive processes. | journal = Jr Neuro Psych and Brain Res: JNPBR | date = June 2019 | url = https://kosmospublishers.com/wp-content/uploads/2019/04/The-Effects-of-Musical-Training-on-Brain-Plasticity-and-Cognitive-Processes.pdf | archive-url = https://web.archive.org/web/20190629032000/https://kosmospublishers.com/wp-content/uploads/2019/04/The-Effects-of-Musical-Training-on-Brain-Plasticity-and-Cognitive-Processes.pdf | archive-date=29 June 2019 }} [143] => [144] => ===In animals=== [145] => {{See also|Brain development|Neural development in humans}} [146] => In a single [[wikt:lifespan|lifespan]], individuals of an animal [[species]] may encounter various changes in brain [[morphology (biology)|morphology]]. Many of these differences are caused by the release of [[hormones]] in the brain; others are the product of [[evolution|evolutionary factors]] or [[Human development (biology)|developmental stages]]. Some changes occur seasonally in species to enhance or generate response behaviors. [147] => [148] => ====Seasonal brain changes==== [149] => Changing brain behavior and morphology to suit other seasonal behaviors is relatively common in animals. These changes can improve the chances of mating during breeding season. Examples of seasonal brain morphology change can be found within many classes and species. [150] => [151] => Within the class [[Aves]], black-capped chickadees experience an increase in the [[volume]] of their [[hippocampus]] and strength of neural connections to the hippocampus during fall months.{{cite journal | vauthors = Barnea A, Nottebohm F | title = Seasonal recruitment of hippocampal neurons in adult free-ranging black-capped chickadees | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 91 | issue = 23 | pages = 11217–11221 | date = November 1994 | pmid = 7972037 | pmc = 45198 | doi = 10.1073/pnas.91.23.11217 | doi-access = free | bibcode = 1994PNAS...9111217B }}{{cite journal | vauthors = Smulders TV, Sasson AD, DeVoogd TJ | title = Seasonal variation in hippocampal volume in a food-storing bird, the black-capped chickadee | journal = Journal of Neurobiology | volume = 27 | issue = 1 | pages = 15–25 | date = May 1995 | pmid = 7643072 | doi = 10.1002/neu.480270103 }} These morphological changes within the hippocampus which are related to [[spatial memory]] are not limited to birds, as they can also be observed in [[rodents]] and [[amphibians]]. In [[songbirds]], many song control nuclei in the brain increase in size during mating season. Among birds, changes in brain morphology to influence song patterns, frequency, and volume are common.{{cite journal | vauthors = Smith GT | title = Seasonal plasticity in the song nuclei of wild rufous-sided towhees | journal = Brain Research | volume = 734 | issue = 1–2 | pages = 79–85 | date = September 1996 | pmid = 8896811 | doi = 10.1016/0006-8993(96)00613-0 | s2cid = 37336866 }} [[Gonadotropin-releasing hormone]] (GnRH) [[immunoreactivity]], or the reception of the hormone, is lowered in [[Common starling|European starlings]] exposed to longer periods of light during the day. [152] => [153] => The [[California sea hare]], a [[gastropod]], has more successful [[inhibitory postsynaptic potential|inhibition]] of egg-laying hormones outside of mating season due to increased effectiveness of inhibitors in the brain. Changes to the inhibitory nature of regions of the brain can also be found in humans and other mammals. In the amphibian [[Bufo japonicus]], part of the [[amygdala]] is larger before breeding and during [[hibernation]] than it is after breeding. [154] => [155] => Seasonal brain variation occurs within many mammals. Part of the hypothalamus of the common [[Sheep|ewe]] is more receptive to GnRH during breeding season than at other times of the year. [[Humans]] experience a change in the "size of the hypothalamic [[suprachiasmatic nucleus]] and [[vasopressin]]-immunoreactive neurons within it" during the fall, when these parts are larger. In the spring, both reduce in size.{{cite journal | vauthors = Tramontin AD, Brenowitz EA | title = Seasonal plasticity in the adult brain | journal = Trends in Neurosciences | volume = 23 | issue = 6 | pages = 251–8 | date = June 2000 | pmid = 10838594 | doi = 10.1016/s0166-2236(00)01558-7 | s2cid = 16888328 }} [156] => [157] => ====Traumatic brain injury research==== [158] => [159] => [[Randy Nudo]]'s group found that if a small [[stroke]] (an infarction) is induced by obstruction of blood flow to a portion of a monkey's motor cortex, the part of the body that responds by movement moves when areas adjacent to the damaged brain area are stimulated. In one study, intracortical microstimulation (ICMS) mapping techniques were used in nine normal monkeys. Some underwent ischemic-infarction procedures and the others, ICMS procedures. The monkeys with ischemic infarctions retained more finger flexion during food retrieval and after several months this deficit returned to preoperative levels. With respect to the distal [[forelimb]] representation, "postinfarction mapping procedures revealed that movement representations underwent reorganization throughout the adjacent, undamaged cortex." Understanding of interaction between the damaged and undamaged areas provides a basis for better treatment plans in stroke patients. Current research includes the tracking of changes that occur in the motor areas of the cerebral cortex as a result of a stroke. Thus, events that occur in the reorganization process of the brain can be ascertained. Nudo is also involved in studying the treatment plans that may enhance recovery from strokes, such as physiotherapy, [[pharmacotherapy]], and electrical-stimulation therapy. [160] => [161] => [[Jon Kaas]], a professor at [[Vanderbilt University]], has been able to show "how somatosensory area 3b and ventroposterior (VP) nucleus of the thalamus are affected by longstanding unilateral dorsal-column lesions at cervical levels in macaque monkeys." Adult brains have the ability to change as a result of injury but the extent of the reorganization depends on the extent of the injury. His recent research focuses on the somatosensory system, which involves a sense of the body and its movements using many senses. Usually, damage of the somatosensory cortex results in impairment of the body perception. Kaas' research project is focused on how these systems (somatosensory, cognitive, motor systems) respond with plastic changes resulting from injury. [162] => [163] => One recent study of neuroplasticity involves work done by a team of doctors and researchers at [[Emory University]], specifically [[Donald Stein]]{{cite web|url=http://www.bme.gatech.edu/facultystaff/faculty_record.php?id=31|archive-url=https://web.archive.org/web/20080624011530/http://www.bme.gatech.edu/facultystaff/faculty_record.php?id=31|archive-date=2008-06-24 |title=Coulter Department of Biomedical Engineering: BME Faculty |publisher=Bme.gatech.edu |access-date=12 June 2010}} and David Wright. This is the first treatment in 40 years that has significant results in treating traumatic brain injuries while also incurring no known side effects and being cheap to administer. Stein noticed that female mice seemed to recover from brain injuries better than male mice, and that at certain points in the [[Estrous cycle|estrus cycle]], females recovered even better. This difference may be attributed to different levels of progesterone, with higher levels of progesterone leading to the faster recovery from brain injury in mice. However, clinical trials showed progesterone offers no significant benefit for traumatic brain injury in human patients.{{Cite web|url=http://news.emory.edu/stories/2014/12/protectiii_results_progesterone_tbi/|title=Progesterone offers no significant benefit in traumatic brain injury clinical trial|date=2014-12-10|website=news.emory.edu|access-date=2016-12-29|archive-date=27 March 2015|archive-url=https://web.archive.org/web/20150327011928/http://news.emory.edu/stories/2014/12/protectiii_results_progesterone_tbi/|url-status=live}} [164] => [165] => ===Aging=== [166] => [167] => [[transcription (biology)|Transcriptional]] profiling of the [[frontal lobe|frontal cortex]] of persons ranging from 26 to 106 years of age defined a set of [[gene]]s with reduced expression after age 40, and especially after age 70. Genes that play central roles in [[synaptic plasticity]] were the most significantly affected by age, generally showing reduced expression over time. There was also a marked increase in cortical [[DNA damage (naturally occurring)|DNA damage]], likely [[DNA oxidation|oxidative DNA damage]], in [[promoter (genetics)|gene promoters]] with aging. [168] => [169] => [[Reactive oxygen species]] appear to have a significant role in the regulation of synaptic plasticity and cognitive function.{{cite journal | vauthors = Massaad CA, Klann E | title = Reactive oxygen species in the regulation of synaptic plasticity and memory | journal = Antioxidants & Redox Signaling | volume = 14 | issue = 10 | pages = 2013–2054 | date = May 2011 | pmid = 20649473 | pmc = 3078504 | doi = 10.1089/ars.2010.3208 }} However age-related increases in reactive oxygen species may also lead to impairments in these functions. [170] => [171] => ===Multilingualism=== [172] => [173] => The beneficial effect of multilingualism on people's behavior and cognition is well-known nowadays. Numerous studies have shown that people who study more than one language have better cognitive functions and flexibilities than people who only speak one language. Bilinguals are found to have longer attention spans, stronger organization and analyzation skills, and a better theory of mind than monolinguals. Researchers have found that the effect of multilingualism on better cognition is due to neuroplasticity. [174] => [175] => In one prominent study, neurolinguists used a [[voxel-based morphometry]] (VBM) method to visualize the structural plasticity of brains in healthy monolinguals and bilinguals. They first investigated the differences in density of grey and white matter between two groups and found the relationship between brain structure and age of language acquisition. The results showed that grey-matter density in the inferior parietal cortex for multilinguals were significantly greater than monolinguals. The researchers also found that early bilinguals had a greater density of grey matter relative to late bilinguals in the same region. The inferior parietal cortex is a brain region highly associated with the language learning, which corresponds to the VBM result of the study.{{cite journal | vauthors = Mechelli A, Crinion JT, Noppeney U, O'Doherty J, Ashburner J, Frackowiak RS, Price CJ | title = Neurolinguistics: structural plasticity in the bilingual brain | journal = Nature | volume = 431 | issue = 7010 | pages = 757 | date = October 2004 | pmid = 15483594 | doi = 10.1038/431757a | hdl-access = free | s2cid = 4338340 | bibcode = 2004Natur.431..757M | hdl = 11858/00-001M-0000-0013-D79B-1 }} [176] => [177] => Recent studies have also found that learning multiple languages not only re-structures the brain but also boosts brain's capacity for plasticity. A recent study found that multilingualism not only affects the grey matter but also white matter of the brain. [[White matter]] is made up of myelinated axons that is greatly associated with learning and communication. Neurolinguists used a [[diffusion tensor imaging]] (DTI) scanning method to determine the white matter intensity between monolinguals and bilinguals. Increased myelinations in white matter tracts were found in bilingual individuals who actively used both languages in everyday life. The demand of handling more than one language requires more efficient connectivity within the brain, which resulted in greater white matter density for multilinguals.{{cite journal | vauthors = Pliatsikas C, Moschopoulou E, Saddy JD | title = The effects of bilingualism on the white matter structure of the brain | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 112 | issue = 5 | pages = 1334–1337 | date = February 2015 | pmid = 25583505 | pmc = 4321232 | doi = 10.1073/pnas.1414183112 | doi-access = free }} [178] => [179] => While it is still debated whether these changes in brain are result of genetic disposition or environmental demands, many evidences suggest that environmental, social experience in early multilinguals affect the structural and functional reorganization in the brain.{{cite journal | vauthors = Draganski B, Gaser C, Busch V, Schuierer G, Bogdahn U, May A | title = Neuroplasticity: changes in grey matter induced by training | journal = Nature | volume = 427 | issue = 6972 | pages = 311–312 | date = January 2004 | pmid = 14737157 | doi = 10.1038/427311a | s2cid = 4421248 |s2cid-access=free | bibcode = 2004Natur.427..311D |url=http://dbm.neuro.uni-jena.de/pdf-files/Draganski-Nature.pdf |url-status=live |archive-url= https://web.archive.org/web/20220626003432/http://dbm.neuro.uni-jena.de/pdf-files/Draganski-Nature.pdf |archive-date= Jun 26, 2022 }}{{cite journal | vauthors = Golestani N, Paus T, Zatorre RJ | title = Anatomical correlates of learning novel speech sounds | journal = Neuron | volume = 35 | issue = 5 | pages = 997–1010 | date = August 2002 | pmid = 12372292 | doi = 10.1016/S0896-6273(02)00862-0 | s2cid = 16089380 |s2cid-access=free | doi-access = free }} [180] => [181] => === Novel treatments of depression === [182] => [183] => Historically, the [[Monoamine hypothesis|monoamine imbalance hypothesis of depression]] played a dominant role in psychiatry and drug development.{{cite journal |author1=Lee, S. |author2=Jeong, J. |author3=Kwak, Y. |author4=Park, S.K. |title=Depression research: where are we now? |journal=Molecular Brain |year=2010 |volume=3 |page=8 |doi=10.1186/1756-6606-3-8 |pmid=20219105 |pmc=2848031 |ref=Mol Brain 3, 8 (2010) |doi-access=free }} However, while traditional [[Antidepressant|antidepressants]] cause a quick increase in [[Norepinephrine|noradrenaline]], [[serotonin]], or [[dopamine]], there is a significant delay in their clinical effect and often an inadequate treatment response.{{cite journal |author1=Rodrigo Machado-Vieira |author2=Jacqueline Baumann |author3=Cristina Wheeler-Castillo |author4=David Latov |author5=Ioline D. Henter |author6=Giacomo Salvadore |author7=Carlos A. Zarate, Jr. |title=The Timing of Antidepressant Effects: A Comparison of Diverse Pharmacological and Somatic Treatments |journal=Pharmaceuticals (Basel, Switzerland) |year=2010 |volume=3 |issue=1 |pages=19–41 |doi=10.3390/ph3010019|pmid=27713241 |pmc=3991019 |doi-access=free }} As neuroscientists pursued this avenue of research, clinical and preclinical data across multiple modalities began to converge on pathways involved in neuroplasticity.{{cite journal |author1=Christopher Pittenger |author2=Ronald S Duman |title=Stress, Depression, and Neuroplasticity: A Convergence of Mechanisms |year=2008 |journal=Neuropsychopharmacology |volume=33 |issue=1 |pages=88–109 |doi=10.1038/sj.npp.1301574 |pmid=17851537 |s2cid=646328 |doi-access=free }} They found a strong inverse relationship between the number of [[synapses]] and severity of depression symptoms{{cite journal |author1=Sophie E. Holmes |author2=Dustin Scheinost |author3=Sjoerd J. Finnema |author4=Mika Naganawa |author5=Margaret T. Davis |author6=Nicole DellaGioia |author7=Nabeel Nabulsi |author8=David Matuskey |author9=Gustavo A. Angarita |author10=Robert H. Pietrzak |author11=Ronald S. Duman |author12=Gerard Sanacora |author13=John H. Krystal |author14=Richard E. Carson |author15=Irina Esterlis |title=Lower synaptic density is associated with depression severity and network alterations |journal=Nature Communications |year=2019 |volume=10 |issue=1 |page=1529 |doi=10.1038/s41467-019-09562-7 |pmid=30948709 |pmc=6449365 |bibcode=2019NatCo..10.1529H }} and discovered that in addition to their [[neurotransmitter]] effect, traditional antidepressants improved neuroplasticity but over a significantly protracted time course of weeks or months.{{cite journal |author1=Ioana Rădulescu |author2=Ana Miruna |author3=Drăgoi Simona |author4=Corina Trifu |author5=Mihai Bogdan Cristea |title=Neuroplasticity and depression: Rewiring the brain's networks through pharmacological therapy |journal=Experimental and Therapeutic Medicine |date=August 5, 2021 |volume=22 |issue=4 |page=1131 |doi=10.3892/etm.2021.10565 |pmid=34504581 |pmc=8383338 }} The search for faster acting antidepressants found success in the pursuit of [[ketamine]], a well-known anesthetic agent, that was found to have potent anti-depressant effects after a single infusion due to its capacity to rapidly increase the number of dendritic spines and to restore aspects of functional connectivity.{{cite journal |author1=Catharine H. Duman |author2=Ronald S. Duman |title=Spine synapse remodeling in the pathophysiology and treatment of depression |journal=Neuroscience Letters |year=2015 |volume=601 |pages=20–29 |doi=10.1016/j.neulet.2015.01.022 |pmid=25582786 |pmc=4497940 }} Additional neuroplasticity promoting compounds with therapeutic effects that were both rapid and enduring have been identified through classes of compounds including [[serotonergic psychedelics]], [[cholinergic]] [[scopolamine]], and other novel compounds. To differentiate between traditional antidepressants focused on monoamine modulation and this new category of fast acting antidepressants that achieve therapeutic effects through neuroplasticity, the term [[psychoplastogen]] was introduced.{{cite web |author1=Calvin Ly |author2=Alexandra C. Greb |author3=Lindsay P. Cameron |author4=Jonathan M. Wong |author5=Eden V. Barragan |author6=Paige C. Wilson |author7=Kyle F. Burbach |author8=Sina Soltanzadeh |author9=Zarandi Alexander Sood |author10=Michael R. Paddy |author11=Whitney C. Duim |author12=Megan Y. Dennis |author13=A. Kimberley McAllister |author14=Kassandra M. Ori-McKenney |author15=John A. Gray |author16=David E. Olson |title=Psychedelics Promote Structural and Functional Neural Plasticity |url=https://www.cell.com/cell-reports/fulltext/S2211-1247(18)30755-1 |website=Cell Reports |access-date=13 July 2022}} [184] => [185] => == See also == [186] => {{col div|colwidth=30em}} [187] => * [[Activity-dependent plasticity]] [188] => * [[Brain training]] [189] => * [[Environmental enrichment (neural)]] [190] => * [[Neural backpropagation]] [191] => * [[Neuronal sprouting]] [192] => * [[Neuroplastic effects of pollution]] [193] => * [[Psychoplastogen]] [194] => * [[Psychedelic drug]] [195] => * [[Kinesiology]] [196] => * [[Spike-timing-dependent plasticity]] [197] => {{div col end}} [198] => [199] => == References == [200] => {{Reflist|refs= [201] => {{cite journal | vauthors = Pascual-Leone A, Freitas C, Oberman L, Horvath JC, Halko M, Eldaief M, Bashir S, Vernet M, Shafi M, Westover B, Vahabzadeh-Hagh AM, Rotenberg A | title = Characterizing brain cortical plasticity and network dynamics across the age-span in health and disease with TMS-EEG and TMS-fMRI | journal = Brain Topography | volume = 24 | issue = 3–4 | pages = 302–315 | date = October 2011 | pmid = 21842407 | pmc = 3374641 | doi = 10.1007/s10548-011-0196-8 }} [202] => [203] => {{cite journal | vauthors = Fuchs E, Flügge G | title = Adult neuroplasticity: more than 40 years of research | journal = Neural Plasticity | volume = 2014 | pages = 541870 | date = 2014 | pmid = 24883212 | pmc = 4026979 | doi = 10.1155/2014/541870 | doi-access = free }} [204] => [205] => {{cite journal | vauthors = Davidson RJ, McEwen BS | title = Social influences on neuroplasticity: stress and interventions to promote well-being | journal = Nature Neuroscience | volume = 15 | issue = 5 | pages = 689–695 | date = April 2012 | pmid = 22534579 | pmc = 3491815 | doi = 10.1038/nn.3093 }} [206] => [207] => {{cite journal | vauthors = Park DC, Huang CM | title = Culture Wires the Brain: A Cognitive Neuroscience Perspective | journal = Perspectives on Psychological Science | volume = 5 | issue = 4 | pages = 391–400 | date = July 2010 | pmid = 22866061 | pmc = 3409833 | doi = 10.1177/1745691610374591 }} [208] => [209] => {{cite journal | vauthors = Shaffer J | title = Neuroplasticity and Clinical Practice: Building Brain Power for Health | journal = Frontiers in Psychology | volume = 7 | pages = 1118 | date = 26 July 2016 | pmid = 27507957 | pmc = 4960264 | doi = 10.3389/fpsyg.2016.01118 | doi-access = free }} [210] => [211] => {{cite journal | vauthors = McEwen BS | title = Redefining neuroendocrinology: Epigenetics of brain-body communication over the life course | journal = Frontiers in Neuroendocrinology | volume = 49 | pages = 8–30 | date = April 2018 | pmid = 29132949 | doi = 10.1016/j.yfrne.2017.11.001 | s2cid = 1681145 | doi-access = }} [212] => [213] => {{cite book |title=Toward a theory of neuroplasticity | veditors = Shaw C, McEachern J |year=2001 |publisher=Psychology Press |location=London, England |isbn=978-1-84169-021-6|title-link=Toward a theory of neuroplasticity}} [214] => [215] => {{cite book | chapter-url = http://psychclassics.yorku.ca/James/Principles/prin4.htm | title = The Principles of Psychology | archive-url = https://web.archive.org/web/20170718003552/http://psychclassics.yorku.ca/James/Principles/prin4.htm | archive-date = 18 July 2017 | vauthors = James W | date = 1890 | chapter = Chapter IV: Habits }} [216] => [217] => {{cite book |title=Synaptic self: how our brains become who we are | vauthors = LeDoux JE |year=2002 |publisher=Viking |location=New York, United States |isbn=978-0-670-03028-6 |page=[https://archive.org/details/synapticselfhowo00ledo/page/137 137] |url=https://archive.org/details/synapticselfhowo00ledo/page/137}} [218] => [219] => [220] => [221] => {{cite journal | vauthors = Ponti G, Peretto P, Bonfanti L | title = Genesis of neuronal and glial progenitors in the cerebellar cortex of peripuberal and adult rabbits | journal = PLOS ONE | volume = 3 | issue = 6 | pages = e2366 | date = June 2008 | pmid = 18523645 | pmc = 2396292 | doi = 10.1371/journal.pone.0002366 | veditors = Reh TA | doi-access = free | bibcode = 2008PLoSO...3.2366P }} [222] => [223] => {{cite journal | vauthors = Wall JT, Xu J, Wang X | title = Human brain plasticity: an emerging view of the multiple substrates and mechanisms that cause cortical changes and related sensory dysfunctions after injuries of sensory inputs from the body | journal = Brain Research. 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}} [271] => *Edelman, Gerald. Bright Air, Brilliant Fire: On the Matter of the Mind (Basic Books, 1992, Reprint edition 1993). {{ISBN|0-465-00764-3}} [272] => *Edelman and Jean-Pierre Changeux, editors, The Brain (Transaction Publishers, 2000). 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Neuroplasticity

Neuroplasticity is the brain's ability to change and reorganize itself throughout an individual's lifespan. This phenomenon allows the brain to adapt to new experiences, learn new skills, and recover from injuries, such as strokes or traumatic brain injuries.

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This phenomenon allows the brain to adapt to new experiences, learn new skills, and recover from injuries, such as strokes or traumatic brain injuries. The concept of neuroplasticity challenges the long-held belief that the brain is a fixed, unchangeable organ. Instead, research has shown that the brain has the ability to form new connections between its neurons, and even generate new neurons in certain regions. The plastic nature of the brain is influenced by various factors, such as genetics, environment, behavior, and age. Understanding neuroplasticity has important implications for fields such as rehabilitation, education, and neuroscience research, as it highlights the potential for individuals to recover from brain damage and develop new abilities throughout their lives.

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