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Neuroplasticity Wikipedia, the free encyclopedia

"Neural plasticity" redirects here. For the journal, see Neural Plasticity (journal).

 For the 2014 album by the band Cold Specks , see  Neuroplasticity (Cold Specks albu)

Contrary to con!entional thouht as e#pressed in this diara, brain functions are not confinedto certain fi#ed locations.

Neuroplasticity , also kno$n as brain plasticity, is an ubrella ter that encopasses bothsynaptic plasticity and non%synaptic plasticity &it refers to chanes in neural path$ays andsynapses due to chanes in beha!ior, en!ironent, neural processes, thinkin, eotions, as $ellas chanes resultin fro bodily injury.' Neuroplasticity has replaced the forerly%held positionthat the brain is a physioloically static oran, and e#plores ho$ % and in $hich $ays % the brainchanes throuhout life.'*

 Neuroplasticity occurs on a !ariety of le!els, ranin fro cellular chanes due to learnin, tolare%scale chanes in!ol!ed in cortical remapping  in response to injury. +he role ofneuroplasticity is $idely reconied in healthy de!elopent, learnin, eory, and reco!eryfro brain daae. -urin ost of the *th century, the scientific consensus aonneuroscientists $as that brain structure is relati!ely iutable after a critical period durin earlychildhood. +his belief has been challened by findins re!ealin that any aspects of the brainreain plastic e!en into adulthood.'/

0ubel and Wiesel had deonstrated that ocular doinance coluns in the lo$est neocortical!isual area, 1, $ere larely iutable after the critical period in de!elopent.'2 Critical periods also $ere studied $ith respect to lanuae3 the resultin data suested that sensory path$ays $ere fi#ed after the critical period. 0o$e!er, studies deterined that en!ironentalchanes could alter beha!ior and conition by odifyin connections bet$een e#istin neuronsand !ia neuroenesis in the hippocapus and other parts of the brain, includin the cerebellu.'4

-ecades of research'5 ha!e no$ sho$n that substantial chanes occur in the lo$est neocortical processin areas, and that these chanes can profoundly alter the pattern of neuronal acti!ation inresponse to e#perience. Neuroscientific research indicates that e#perience can actually chane

 both the brain6s physical structure (anatoy) and functional oraniation ( physioloy).

 Neuroscientists are currently enaed in a reconciliation of critical period studies deonstratinthe iutability of the brain after de!elopent $ith the ore recent research sho$in ho$ the

 brain can, and does, chane in response to hitherto unsuspected stiuli.'7'8

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Contents

• Neurobioloy 

o . Cortical aps

• * 9pplications and e#aple 

o *. +reatent of brain daae

o *.* 1ision

o *./ +reatent of learnin difficulties

o *.2 -urin operation of brain%achine interfaces

o *.4 Sensory prostheses

o *.5 Phanto libs

o *.7 Chronic pain

o *.8 :editation

o *.; Fitness and e#ercise

o *. 0uan echolocation

o *. <n anials 

*.. Seasonal brain chanes

• / =tyoloy

• 2 0istory 

o 2. Proposal

o 2.* >esearch and disco!ery

• 4 See also

• 5 >eferences

• 7 Further readin

•

8 =#ternal links

Neurobiology

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?ne of the fundaental principles of ho$ neuroplasticity functions is linked to the concept ofsynaptic prunin, the idea that indi!idual connections $ithin the brain are constantly beinreo!ed or recreated, larely dependent upon ho$ they are used. +his concept is captured in theaphoris, "neurons that fire toether, $ire toether"@"neurons that fire apart, $ire apart,"suariin 0ebbian theory. <f there are t$o nearby neurons that often produce an ipulsesiultaneously, their cortical aps ay becoe one. +his idea also $orks in the opposite $ay,

i.e. that neurons $hich do not reularly produce siultaneous ipulses $ill for different aps.

Cortical maps

Cortical oraniation, especially for the sensory systes, is often described in ters of aps.'; For e#aple, sensory inforation fro the foot, projects to one cortical site and the projectionsfro the hand, taret another site. 9s the result of this soatotopic oraniation of sensoryinputs to the corte#, cortical representation of the body resebles a ap (or hounculus).

<n the late ;7s and early ;8s, se!eral roups bean e#plorin the ipacts of reo!in portions of the sensory inputs. :ichael :erenich, Aon Baas and -ou >asusson used the

cortical ap as their dependent !ariable. +hey found&and this has been since corroborated by a$ide rane of labs&that if the cortical ap is depri!ed of its input it $ill becoe acti!ated at alater tie in response to other, usually adjacent inputs. :erenichs (;82) study in!ol!ed theappin of o$l onkey hands before and after aputation of the third diit. Defore aputation,there $ere fi!e distinct areas, one correspondin to each diit of the e#periental hand. Si#ty%t$o days follo$in aputation of the third diit, the area in the cortical ap forerly occupied

 by that diit had been in!aded by the pre!iously adjacent second and fourth diit ones. +heareas representin diit one and fi!e are not located directly beside the area representin diitthree, so these reions reained, for the ost part, unchaned follo$in aputation.' +hisstudy deonstrates that only those reions borderin a certain area $ill in!ade it to alter thecortical ap. <n the soatic sensory syste, in $hich this phenoenon has been ostthorouhly in!estiated, A+ Wall and A Eu ha!e traced the echaniss underlyin this plasticity.>e%oraniation is not cortically eerent, but occurs at e!ery le!el in the processin hierarchy3this produces the ap chanes obser!ed in the cerebral corte#.'

:erenich and Willia Aenkins (;;) initiated studies relatin sensory e#perience, $ithout patholoical perturbation, to cortically obser!ed plasticity in the priate soatosensory syste, $ith the findin that sensory sites acti!ated in an attended operant beha!ior  increase in theircortical representation. Shortly thereafter, Ford =bner and colleaues (;;2) ade siilar effortsin the rodent $hisker   barrel corte# (also soatic sensory syste). +hese t$o roups larelydi!ered o!er the years. +he rodent $hisker barrel efforts becae a focus for =bner, :atthe$

-iaond, :ichael 9rstron%Aaes, >obert Sachde!, and Be!in Fo#. reat inroads $ere adein identifyin the locus of chane as bein at cortical synapses e#pressin  N:-9 receptors, andin iplicatin cholineric inputs as necessary for noral e#pression. 0o$e!er, the rodent studies$ere poorly focused on the beha!ioral end, and >on Frosti and -aniel Polley (;;;, *2)identified beha!ioral anipulations as causin a substantial ipact on the cortical plasticity inthat syste.

:erenich and -+ Dlake (**, *4, *5) $ent on to use cortical iplants to study thee!olution of plasticity in both the soatosensory and auditory systes. Doth systes sho$siilar chanes $ith respect to beha!ior . When a stiulus is coniti!ely associated $ithreinforceent, its cortical representation is strenthened and enlared. <n soe cases, cortical

representations can increase t$o to threefold in G* days at the tie at $hich a ne$ sensoryotor beha!ior is first acHuired, and chanes are larely finished $ithin at ost a fe$ $eeks.Control studies sho$ that these chanes are not caused by sensory e#perience aloneI they reHuire

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learnin about the sensory e#perience, and are stronest for the stiuli that are associated $ithre$ard, and occur $ith eHual ease in operant and classical conditionin beha!iors.

9n interestin phenoenon in!ol!in cortical aps is the incidence of phanto libs. Phantolibs are e#perienced by people $ho ha!e underone aputations in hands, ars, and les, butit is not liited to e#treities. 9lthouh the neuroloical basis of phanto libs is still not

entirely understood it is belie!ed that cortical reoraniation plays an iportant role.'*

 Noran -oide, follo$in the lead of :ichael :erenich, separates anifestations ofneuroplasticity into adaptations that ha!e positi!e or neati!e beha!ioral conseHuences. Fore#aple, if an oranis can reco!er after a stroke to noral le!els of perforance, thatadapti!eness could be considered an e#aple of "positi!e plasticity". Chanes such as ane#cessi!e le!el of neuronal ro$th leadin to spasticity or tonic paralysis, or an e#cessi!e releaseof neurotransitters in response to injury $hich could kill ner!e cells, $ould ha!e to beconsidered "neati!e" plasticity. <n addition, dru addiction and obsessi!e%copulsi!e disorderare deeed e#aples of "neati!e plasticity" by -r. -oide, as the synaptic re$irin resultin inthese beha!iors is also hihly aladapti!e.'*'/

9 *4 study found that the effects of neuroplasticity occur e!en ore rapidly than pre!iouslye#pected. :edical students6 brains $ere iaed durin the period $hen they $ere studyin fortheir e#as. <n a atter of onths, the students6 ray atter increased sinificantly in the

 posterior and lateral parietal corte#.'2

Applications and example

Treatment of brain damage

9 surprisin conseHuence of neuroplasticity is that the brain acti!ity associated $ith a i!enfunction can o!e to a different location3 this can result fro noral e#perience and also occursin the process of reco!ery fro brain injury. Neuroplasticity is the fundaental issue thatsupports the scientific basis for treatent of acHuired brain injury $ith oal%directed e#perientialtherapeutic proras in the conte#t of rehabilitation approaches to the functional conseHuencesof the injury.

+he adult brain is not entirely "hard%$ired" $ith fi#ed neuronal circuits. +here are anyinstances of cortical and subcortical re$irin of neuronal circuits in response to trainin as $ellas in response to injury. +here is solid e!idence that neuroenesis (birth of brain cells) occurs inthe adult, aalian brain&and such chanes can persist $ell into old ae. '/ +he e!idence for

neuroenesis is ainly restricted to the hippocapus and olfactory bulb, but current research hasre!ealed that other parts of the brain, includin the cerebellu, ay be in!ol!ed as $ell. '4

<n the rest of the brain, neurons can die, but they cannot be created. 0o$e!er, there is no$ aplee!idence for the acti!e, e#perience%dependent re%oraniation of the synaptic net$orks of the

 brain in!ol!in ultiple inter%related structures includin the cerebral corte#. +he specific detailsof ho$ this process occurs at the olecular and ultrastructural le!els are topics of acti!eneuroscience research. +he anner in $hich e#perience can influence the synaptic oraniationof the brain is also the basis for a nuber of theories of brain function includin the eneraltheory of ind and episteoloy referred to as Neural -ar$inis and de!eloped by

iunoloist Nobel laureate erald =delan. +he concept of neuroplasticity is also central totheories of eory and learnin that are associated $ith e#perience%dri!en alteration of synapticstructure and function in studies of classical conditionin in in!ertebrate anial odels such as9plysia. +his latter prora of neuroscience research has follo$ed fro the round%breakin

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$ork of another Nobel laureate, =ric Bandel, and his colleaues at Colubia Jni!ersity Colleeof Physicians and Sureons.

Paul Dach%y%>ita, deceased in *5, $as the "father of sensory substitution and brain plasticity."'4 <n $orkin $ith a patient $hose !estibular syste had been daaed he de!elopedDrainPort,'5 a achine that "replaces her !estibular apparatus and '$ill send balance sinals to

her brain fro her tonue."'*

 9fter she had used this achine for soe tie it $as no lonernecessary, as she reained the ability to function norally. 0er balancin%act days $ere o!er.'7

Plasticity is the ajor e#planation for this phenoenon. Decause her !estibular syste $as"disoranied" and sendin rando rather than coherent sinals, the apparatus found ne$

 path$ays around the daaed or blocked neural path$ays, helpin to reinforce the sinals that$ere sent by reainin healthy tissues. Dach%y%>ita e#plained plasticity by sayin, "<f you aredri!in fro here to :il$aukee and the ain bride oes out, first you are paralyed. +hen youtake old secondary roads throuh the farland. +hen you use these roads ore3 you find shorter

 paths to use to et $here you $ant to o, and you start to et there faster. +hese "secondary"neural path$ays are "unasked" or e#posed and strenthened as they are used. +he "unaskin"

 process is enerally thouht to be one of the principal $ays in $hich the plastic brain reoraniesitself."'*

>andy Nudo6s roup found that if a sall stroke (an infarction) is induced by obstruction of blood flo$ to a portion of a onkeys otor corte#, the part of the body that responds byo!eent $ill o!e $hen areas adjacent to the daaed brain area are stiulated. <n one study,intracortical icrostiulation (<C:S) appin techniHues $ere used in nine noral onkeys.Soe under$ent ischeic%infarction procedures and the others, <C:S procedures. +he onkeys$ith ischeic infarctions retained ore finer fle#ion durin food retrie!al and after se!eralonths this deficit returned to preoperati!e le!els.'8 With respect to the distal forelibrepresentation, "postinfarction appin procedures re!ealed that o!eent representationsunder$ent reoraniation throuhout the adjacent, undaaed corte#."'8 Jnderstandin ofinteraction bet$een the daaed and undaaed areas pro!ides a basis for better treatent

 plans in stroke patients. Current research includes the trackin of chanes that occur in the otor areas of the cerebral corte# as a result of a stroke. +hus, e!ents that occur in the reoraniation

 process of the brain can be ascertained. Nudo is also in!ol!ed in studyin the treatent plansthat ay enhance reco!ery fro strokes, such as physiotherapy, pharacotherapy, and electrical%stiulation therapy.

 Neuroplasticity is ainin popularity as a theory that, at least in part, e#plains ipro!eents infunctional outcoes $ith physical therapy post stroke. >ehabilitation techniHues that ha!e

e!idence to suest cortical reoraniation as the echanis of chane include Constraint%induced o!eent therapy, functional electrical stiulation, treadill trainin $ith body%$eihtsupport, and !irtual reality therapy. >obot assisted therapy is an eerin techniHue, $hich isalso hypothesied to $ork by $ay of neuroplasticity, thouh there is currently insufficiente!idence to deterine the e#act echaniss of chane $hen usin this ethod.';

Aon Baas, a professor at 1anderbilt Jni!ersity, has been able to sho$ "ho$ soatosensory area/b and !entroposterior (1P) nucleus of the thalaus are affected by lonstandin unilateraldorsal%colun lesions at cer!ical le!els in acaHue onkeys." '* 9dult brains ha!e the ability tochane as a result of injury but the e#tent of the reoraniation depends on the e#tent of theinjury. 0is recent research focuses on the soatosensory syste, $hich in!ol!es a sense of the

 body and its o!eents usin any senses. Jsually $hen people daae the soatosensorycorte#, ipairent of the body perceptions are e#perienced. 0e is tryin to see ho$ thesesystes (soatosensory, coniti!e, otor systes) are plastic as a result of injury.'*

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?ne of the ost recent applications of neuroplasticity in!ol!es $ork done by a tea of doctorsand researchers at =ory Jni!ersity, specifically -r. -onald Stein ($ho has been in the field for o!er three decades)'* and -r. -a!id Wriht. +his is the first treatent in 2 years that hassinificant results in treatin trauatic brain injuries $hile also incurrin no kno$n side effectsand bein cheap to adinister .'** -r. Stein noticed that feale ice seeed to reco!er fro

 brain injuries better than ale ice. 9lso in feales, he noticed that at certain points in the

estrus cycle feales reco!ered e!en ore. 9fter lots of research, they attributed this differencedue to the le!els of proesterone. +he hihest le!el of proesterone present led to the fastestreco!ery of brain injury in these ice.

+hey de!eloped a treatent that includes increased le!els of proesterone injections to i!e to brain%injured patients. "9dinistration of proesterone after trauatic brain injury'*/ (+D<) andstroke reduces edea, inflaation, and neuronal cell death, and enhance spatial referenceeory and sensory otor reco!ery."'*2 <n their clinical trials, they had a roup of se!erelyinjured patients that after the three days of proesterone injections had a 5K reduction inortality.'** SaL $as in a horrific car accident that left hi $ith arinal brain acti!ity3accordin to the doctors, he $as one point a$ay fro bein brain dead. 0is parents decided to

ha!e hi participate in -r. Steins clinical trial and he $as i!en the three%day proesteronetreatent. +hree years after the accident, he had achie!ed an inspirin reco!ery $ith no braincoplications and the ability to li!e a healthy, noral life.'**

Stein has done soe studies in $hich beneficial effects ha!e been seen to be siilar in aed ratsto those seen in youthful rats. 9s there are physioloical differences in the t$o ae roups, theodel $as t$eaked for the elderly anials by reducin their stress le!els $ith increased physicalcontact. -urin surery, anesthesia $as kept at a hiher o#yen le!el $ith lo$er o!erallisoflurane percentae and "the aed anials $ere i!en subcutaneous lactated riners solution 

 post%surery to replace fluids lost throuh increased bleedin."'*4 +he proisin results of proesterone treatents "could ha!e a sinificant ipact on the clinical anaeent of +D<."'*4 +hese treatents ha!e been sho$n to $ork on huan patients $ho recei!e treatent soon afterthe +D<. 0o$e!er, -r. Stein no$ focuses his research on those persons $ho ha!e lonstandintrauatic brain injury in order to deterine if proesterone treatents $ill assist the in thereco!ery of lost functions as $ell.

9 study published in the New England ournal o! "edicine in *2 detailin the results of aulti%center N<0%funded phase <<< clinical trial of 88* patients found that treatent of acutetrauatic brain injury $ith the horone proesterone pro!ides no sinificant benefit to patients$hen copared $ith placebo. '*5

Vision

9fter decades in $hich the assuption that binocular !ision, in particular stereopsis, had to beachie!ed in early childhood lest it could ne!er be ained, in recent years the successfulipro!eents in persons $ith ablyopia, con!erence insufficiency or stereo !ision anoaliesha!e becoe prie e#aples of neuroplasticity3 binocular !ision ipro!eents and stereopsisreco!ery are no$ acti!e areas of scientific and clinical research. '*7'*8'*;

Treatment of learning difficulties

:ichael :erenich de!eloped a series of "plasticity%based coputer proras kno$n as Fast

ForWord."'citation needed  FastForWord offers se!en brain e#ercises to help $ith the lanuae andlearnin deficits of dysle#ia. <n a recent study, e#periental trainin $as done in adults to see ifit $ould help to counteract the neati!e plasticity that results fro ae%related coniti!e decline

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(9>C-). +he =+ desin included si# e#ercises desined to re!erse the dysfunctions caused by9>C- in conition, eory, otor control, and so on. 9fter use of the =+ prora for 8G$eeks, there $as a "sinificant increase in task%specific perforance." 'citation needed  +he datacollected fro the study indicated that a neuroplasticity%based prora could notably ipro!econiti!e function and eory in adults $ith 9>C-.

During operation of brain-machine interfaces

Drain%achine interface (D:<) is a rapidly de!elopin field of  neuroscience. 9ccordin to theresults obtained by :ikhail Mebede!, :iuel Nicolelis and their colleaues,'/ operation of D:<sresults in incorporation of artificial actuators into brain representations. +he scientists sho$edthat odifications in neuronal representation of the onkey6s hand and the actuator that $ascontrolled by the onkey brain occurred in ultiple cortical areas $hile the onkey operated aD:<. <n these sinle day e#perients, onkeys initially o!ed the actuator by pushin a

 joystick. 9fter appin out the otor neuron ensebles, control of the actuator $as s$itched tothe odel of the ensebles so that the brain acti!ity, and not the hand, directly controlled theactuator. +he acti!ity of indi!idual neurons and neuronal populations becae less representati!e

of the anial6s hand o!eents $hile representin the o!eents of the actuator. Presuablyas a result of this adaptation, the anials could e!entually stop o!in their hands yet continueto operate the actuator. +hus, durin D:< control, cortical ensebles plastically adapt, $ithintens of inutes, to represent beha!iorally sinificant otor paraeters, e!en if these are notassociated $ith o!eents of the anial6s o$n lib.

9cti!e laboratory roups include those of Aohn -onohue at Dro$n, >ichard 9ndersen atCaltech, Brishna Shenoy at Stanford, Nicholas 0atsopoulos of Jni!ersity of Chicao, 9ndySch$art at Jni!ersity of Pittsburh, Sandro :ussa%<!aldi at North$estern and :iuel Nicolelisat -uke. -onohue and Nicolelis6 roups ha!e independently sho$n that anials can controle#ternal interfaces in tasks reHuirin feedback, $ith odels based on acti!ity of cortical neurons,and that anials can adapti!ely chane their inds to ake the odels $ork better. -onohue6sroup took the iplants fro >ichard Norann6s lab at Jtah (the "Jtah" array), and ipro!ed it

 by chanin the insulation fro polyiide to parylene%c, and coercialied it throuh thecopany Cyberkinetics. +hese efforts are the leadin candidate for the first huan trials on a

 broad scale for otor cortical iplants to help Huadripleic or locked%in patients counicate$ith the outside $orld.

Sensory prostheses

 Neuroplasticity is in!ol!ed in the de!elopent of sensory function. +he brain is born iature

and it adapts to sensory inputs after birth. <n the auditory syste, conenital hearin ipairent,a rather freHuent inborn condition affectin of ne$borns, has been sho$n to affectauditory de!elopent, and iplantation of a sensory prostheses acti!atin the auditory systehas pre!ented the deficits and induced functional aturation of the auditory syste '/ -ue to asensiti!e period for plasticity, there is also a sensiti!e period for such inter!ention $ithin the first*G2 years of life. ConseHuently, in prelinually deaf children, early cochlear iplantation as arule allo$s to learn other lanuae and acHuire acoustic counication.'/*

Phantom limbs

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9 diaraatic e#planation of the irror bo#. +he patient places the ood lib into one side ofthe bo# (in this case the riht hand) and the aputated lib into the other side. -ue to the irror,the patient sees a reflection of the ood hand $here the issin lib $ould be (indicated inlo$er contrast). +he patient thus recei!es artificial !isual feedback that the "resurrected" lib is

no$ o!in $hen they o!e the ood hand.:ain articlesI Phanto lib and :irror bo#

+he e#perience of Phanto libs is a phenoenon in $hich a person continues to feel pain orsensation $ithin a part of their body $hich has been aputated. +his is stranely coon,occurrin in 5%8K of aputees.'// 9n e#planation for this refers to the concept ofneuroplasticity, as the cortical aps of the reo!ed libs are belie!ed to ha!e becoe enaed$ith the area around the in the postcentral yrus. +his results in acti!ity $ithin the surroundinarea of the corte# bein isinterpreted by the area of the corte# forerly responsible for theaputated lib.

+he relationship bet$een phanto libs and neuroplasticity is a cople# one. <n the early ;;s1.S. >aachandran theoried that phanto libs $ere the result of cortical reappin.0o$e!er, in ;;4 0erta Flor and her colleaues deonstrated that cortical reappin occursonly in patients $ho ha!e phanto pain.'/2 0er research sho$ed that phanto lib pain (ratherthan referred sensations) $as the perceptual correlate of cortical reoraniation.'/4 +his

 phenoenon is soeties referred to as aladapti!e plasticity.

<n *; Morier :oseley and Peter Druer carried out a rearkable e#perient in $hich theyencouraed ar aputee subjects to use !isual iaery to contort their phanto libs intoipossible confiurations. Four of the se!en subjects succeeded in perforin ipossible

o!eents of the phanto lib. +his e#perient suests that the subjects had odified theneural representation of their phanto libs and enerated the otor coands needed toe#ecute ipossible o!eents in the absence of feedback fro the body.'/5 +he authors statedthatI"<n fact, this findin e#tends our understandin of the brain6s plasticity because it is e!idencethat profound chanes in the ental representation of the body can be induced purely by internal

 brain echaniss%%the brain truly does chane itself."

Chronic pain

:ain articleI Chronic pain

<ndi!iduals $ho suffer fro chronic pain e#perience proloned pain at sites that ay ha!e been pre!iously injured, yet are other$ise currently healthy. +his phenoenon is related toneuroplasticity due to a aladapti!e reoraniation of the ner!ous syste, both peripherally and

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centrally. -urin the period of tissue daae, no#ious stiuli and inflaation cause anele!ation of nocicepti!e input fro the periphery to the central ner!ous syste. Prolonednociception fro periphery $ill then elicit a neuroplastic response at the cortical le!el to chaneits soatotopic oraniation for the painful site, inducin central sensitiation.'/7 For instance,indi!iduals e#periencin cople# reional pain syndroe deonstrate a diinished corticalsoatotopic representation of the hand contralaterally as $ell as a decreased spacin bet$een the

hand and the outh.'/8

 9dditionally, chronic pain has been reported to sinificantly reduce the!olue of rey atter  in the brain lobally, and ore specifically at the prefrontal corte# andriht thalaus.'/; 0o$e!er, follo$in treatent, these abnoralities in cortical reoraniationand rey atter !olue are resol!ed, as $ell as their syptos. Siilar results ha!e beenreported for phanto lib pain,'2 chronic lo$ back pain'2 and carpal tunnel syndroe.'2*

Meditation

:ain articleI >esearch on editation

9 nuber of studies ha!e linked editation practice to differences in cortical thickness or

density of ray atter.'2/'22'24 ?ne of the ost $ell%kno$n studies to deonstrate this $as led by Sara Maar , fro 0ar!ard Jni!ersity, in *.'25 >ichard -a!idson, a neuroscientist at theJni!ersity of Wisconsin, has led e#perients in cooperation $ith the -alai Maa on effects ofeditation on the brain. 0is results suest that lon%ter, or short%ter practice of editationresults in different le!els of acti!ity in brain reions associated $ith such Hualities as attention,an#iety, depression, fear , aner , the ability of the body to heal itself, and so on. +hese functionalchanes ay be caused by chanes in the physical structure of the brain. '27'28'2;'4

itness and exercise

:ain articleI Neurobioloical effects of physical e#ercise Neuroplasticity and neuroenesis

<n a *; study, scientists ade t$o roups of ice s$i a $ater ae, and then in a separatetrial subjected the to an unpleasant stiulus to see ho$ Huickly they $ould learn to o!e a$ayfro it. +hen, o!er the ne#t four $eeks they allo$ed one roup of ice to run inside their rodent$heels, an acti!ity ost ice enjoy, $hile they forced the other roup to $ork harder oninitreadills at a speed and duration controlled by the scientists. +hey then tested both roupsaain to track their learnin skills and eory. Doth roups of ice ipro!ed their

 perforances in the $ater ae fro the earlier trial. Dut only the e#tra%$orked treadillrunners $ere better in the a!oidance task, a skill that, accordin to neuroscientists, deands aore coplicated coniti!e response.'4

+he ice $ho $ere forced to run on the treadills sho$ed e!idence of olecular chanes inse!eral portions of their brains $hen !ie$ed under a icroscope, $hile the !oluntary $heel%runners had chanes in only one area. "?ur results support the notion that different fors ofe#ercise induce neuroplasticity chanes in different brain reions," Chauyin A. Aen, a professorof physioloy and an author of the study, said. '4* Siilar results ha!e ean$hile been found forhuans. '8

!uman echolocation

0uan echolocation is a learned ability for huans to sense their en!ironent fro echoes. +hisability is used by soe blind people to na!iate their en!ironent and sense their surroundinsin detail. Studies in * '4/ and * '42 usin Functional anetic resonance iain techniHues ha!e sho$n that parts of the brain associated $ith !isual processin are adapted for

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the ne$ skill of echolocation. Studies $ith blind patients, for e#aple, suest that the click%echoes heard by these patients $ere processed by brain reions de!oted to !ision rather thanaudition.'44

"n animals

See alsoI Drain de!elopent and Neural de!elopent in huans

<n a sinle lifespan, indi!iduals in an anial species ay encounter !arious chanes in brainorpholoy. :any of these differences are caused by the release of horones in the brain3others are the product of e!olutionary factors or de!elopental staes.'45'47'48'4; Soe chanesoccur seasonally in species to enhance or enerate response beha!iors.

Seasonal brain changes

Chanin brain beha!ior and orpholoy to suit other seasonal beha!iors is relati!ely coonin anials.'5 +hese chanes can ipro!e the chances of atin durin breedin season.'45'47'48

'5'5'5* =#aples of seasonal brain orpholoy chane can be found $ithin any classes andspecies.

Within the class 9!es, black%capped chickadees e#perience an increase in the !olue of thehippocapus and strenth of neural connections to the hippocapus durin fall onths.'5/'52 +his chane in brain orpholoy for spatial eory $ithin the hippocapus is not liited to

 birds, and affects soe rodents and aphibians.'5 <n sonbirds, any son control nuclei in the brain increase in sie durin atin season.'5 9on birds, chanes in brain orpholoy toinfluence son patterns, freHuency, and !olue are coon.'54 onadotropin%releasin horone(n>0) iunoreacti!ity, or the reception of the horone, is lo$ered in =uropean starlins 

e#posed to loner periods of liht durin the day.

'45'47

+he California sea hare, a astropod, has ore successful inhibition of e%layin horonesoutside of atin season due to increased effecti!eness of inhibitors in the brain. '48 Chanes tothe inhibitory nature of reions of the brain can also be found in huans and other aals. '4; <n the aphibian Dufo japonicus, part of the aydala is larer before breedin and durinhibernation than it is after breedin.'5

Seasonal brain !ariation occurs $ithin any aals. Part of the hypothalaus of the coone$e is ore recepti!e to n>0 durin breedin season than at other ties of the year .'5* 0uans e#perience a chane in the "sie of the hypothalaic suprachiasatic nucleus and

!assopressin%iunoreacti!e neurons $ithin it"'4;

 durin the fall, $hen these parts are larer. <nthe sprin, both reduce in sie.'55

#tymology

Plasticity $as first applied to beha!ior in 8; by Willia Aaes in #he $rinciples o!

 $sychology,'57 thouh the idea $as larely nelected for the ne#t fifty years'citation needed . +he first person to use the ter neural plasticity appears to ha!e been the Polish neuroscientist AeryBonorski.'58

i!en the central iportance of neuroplasticity, an outsider $ould be fori!en for assuin thatit $as $ell defined and that a basic and uni!ersal frae$ork ser!ed to direct current and futurehypotheses and e#perientation. Sadly, ho$e!er, this is not the case. While any neuroscientists

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use the $ord neuroplasticity as an ubrella ter it eans different thins to different researchersin different subfields ... <n brief, a utually areed upon frae$ork does not appear to e#ist. '5;

!istory

Proposal

Jntil around the ;7s, an accepted idea across neuroscience $as that the ner!ous syste $asessentially fi#ed throuhout adulthood, both in ters of brain functions, as $ell as the idea that it$as ipossible for ne$ neurons to de!elop after birth.'7

<n 7;/, <talian anatoist :ichele 1iceno :alacarne described e#perients in $hich he pairedanials, trained one of the pair e#tensi!ely for years, and then dissected both. 0e disco!ered thatthe cerebellus of the trained anials $ere substantially larer. Dut, these findins $eree!entually forotten.'7 +he idea that the brain and its functions are not fi#ed throuhoutadulthood $as proposed in 8; by Willia Aaes in #he $rinciples o! $sychology, thouh the

idea $as larely nelected.'57

$esearch and disco%ery

<n ;*/, Barl Mashley conducted e#perients on rhesus onkeys $hich deonstrated chanesin neuronal path$ays, $hich he concluded to be e!idence of plasticity, althouh despite this, as$ell as further e#aples of research suestin this, the idea of neuroplasticity $as not $idelyaccepted by neuroscientists. 0o$e!er, ore sinificant e!idence bean to be produced in the;5s and after, notably fro scientists includin Paul Dach%y%>ita, :ichael :erenich alon$ith Aon Baas, as $ell as se!eral others.'7'7*

<n the ;5s, Paul Dach%y%>ita in!ented a de!ice that allo$ed blind people to read, percei!eshado$s, and distinuish bet$een close and distant objects. +his "achine $as one of the firstand boldest applications of neuroplasticity."'* +he patient sat in an electrically stiulated chairthat had a lare caera behind it $hich scanned the area, sendin electrical sinals of the iaeto four hundred !ibratin stiulators on the chair aainst the patients skin. +he si# subjects ofthe e#perient $ere e!entually able to reconie a picture of the superodel +$iy.'*

<t ust be ephasied that these people $ere conenitally blind and had pre!iously not beenable to see. Dach%y%>ita belie!ed in sensory substitution3 if one sense is daaed, your othersenses can soeties take o!er. 0e thouht the skin and its touch receptors could act as a retina(usin one sense for another '7/). <n order for the brain to interpret tactile inforation and con!ertit into !isual inforation, it has to learn soethin ne$ and adapt to the ne$ sinals. +he brain6scapacity to adapt iplied that it possessed plasticity. 0e thouht, "We see $ith our brains, not$ith our eyes."'*

9 traic stroke that left his father paralyed inspired Dach%y%>ita to study brain rehabilitation.0is brother, a physician, $orked tirelessly to de!elop therapeutic easures $hich $ere sosuccessful that the father reco!ered coplete functionality by ae 58 and $as able to li!e anoral, acti!e life $hich e!en included ountain clibin. "0is fathers story $as firsthande!idence that a Olate reco!ery could occur e!en $ith a assi!e lesion in an elderly person."'* 0e found ore e!idence of this possible brain reoraniation $ith Shepherd <!ory Fran6s $ork.'72

 ?ne study in!ol!ed stroke patients $ho $ere able to reco!er throuh the use of brainstiulatin e#ercises after ha!in been paralyed for years. "Fran understood the iportance ofinterestin, oti!atin rehabilitationI OJnder conditions of interest, such as that of copetition,

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the resultin o!eent ay be uch ore efficiently carried out than in the dull, routinetrainin in the laboratory(Fran, ;*, p.;/)."'74 +his notion has led to oti!ationalrehabilitation proras that are used today.

=leanor :auire docuented chanes in hippocapal structure associated $ith acHuirin thekno$lede of Mondons layout in local ta#i dri!ers.'75'77'78 9 redistribution of rey atter $as

indicated in Mondon +a#i -ri!ers copared to controls. +his $ork on hippocapal plasticity notonly interested scientists, but also enaed the public and edia $orld%$ide.

:ichael :erenich is a neuroscientist $ho has been one of the pioneers of neuroplasticity foro!er three decades. 0e has ade soe of "the ost abitious clais for the field % that braine#ercises ay be as useful as drus to treat diseases as se!ere as schiophrenia % that plasticitye#ists fro cradle to the ra!e, and that radical ipro!eents in coniti!e functionin % ho$ $elearn, think, percei!e, and reeber are possible e!en in the elderly." '* :erenichs $ork $asaffected by a crucial disco!ery ade by -a!id 0ubel and +orsten Wiesel in their $ork $ithkittens. +he e#perient in!ol!ed se$in one eye shut and recordin the cortical brain aps.0ubel and Wiesel sa$ that the portion of the kittens brain associated $ith the shut eye $as not

idle, as e#pected. <nstead, it processed !isual inforation fro the open eye. <t $as "as thouhthe brain didnt $ant to $aste any Ocortical real estate and had found a $ay to re$ire itself." '*

+his iplied neuroplasticity durin the critical period. 0o$e!er, :erenich arued thatneuroplasticity could occur beyond the critical period. 0is first encounter $ith adult plasticitycae $hen he $as enaed in a postdoctoral study $ith Clinton Woosley. +he e#perient $as

 based on obser!ation of $hat occurred in the brain $hen one peripheral ner!e $as cut andsubseHuently reenerated. +he t$o scientists icroapped the hand aps of onkey brains

 before and after cuttin a peripheral ner!e and se$in the ends toether. 9fter$ards, the handap in the brain that $as e#pected to be jubled $as nearly noral. +his $as a substantial

 breakthrouh. :erenich asserted that "if the brain ap could noralie its structure in responseto abnoral input, the pre!ailin !ie$ that $e are born $ith a hard$ired syste had to be$ron. +he brain had to be plastic."'*

See also

• 9cti!ity%dependent plasticity

• =n!ironental enrichent (neural)

•  Neuroplastic effects of pollution

• Binesioloy

$eferences

.

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 brain%deri!ed neurotrophic factor and synaptotain <.". ournal o! $hysiology .'/ (/)I /**G /*/. doiI./@jphysiol.*;.7/88.    retchen >eynolds (5 Septeber *;). "Phys =dI What Sort of =#ercise Can :ake QouSarterX".  Ne$ Qork +ies.     0uan =cholocation, Aournal of 1ision / 9uust * !ol. no. 7 article 4httpI@@$$$.journalof!ision.or@content@@7@4.abstract   

  Neural Correlates of Natural 0uan =cholocation in =arly and Mate Dlind =cholocation=#perts,PMoS ?ne, *4 :ay *, doiI./7@journal.pone.*5* , 

httpI@@$$$.plosone.or@article@infoIdoiK*F./7K*Fjournal.pone.*5*      +haler, M3 9rnot, S.>.3 oodale, :.9 (*). "Neural correlates of natural huanecholocation in early and late blind echolocation e#perts". $ublic (ibrary o! Science * (4).    -.:. Parry et al. <unocytocheical localiation of n>0 precursor in the hypothalausof =uropean starlins durin se#ual aturation and photorefractoriness A. Neuroendocrinol., ;(;;7), pp. */4G*2/    -.:. Parry, 9.>. oldsith Jltrastructural e!idence for chanes in synaptic input to thehypothalaic luteiniin horone%releasin horone neurons in photosensiti!e and

 photorefractory starlins A. Neuroendocrinol., 4 (;;/), pp. /87G/;4

 N.M. Wayne et al. Seasonal fluctuations in the secretory response of neuroendocrine cells of9plysia californica to inhibitors of protein kinase 9 and protein kinase C en. Cop.=ndocrinol., ; (;;8), pp. /45G/54

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    :.9. 0ofan, -.F. S$aab Seasonal chanes in the suprachiasatic nucleus of an Neurosci. Mett., /; (;;*), pp. *47G*5    F. Notteboh 9 brain for all seasonsI cyclical anatoical chanes in son control nuclei ofthe canary brain Science, *2 (;8), pp. /58G/7    S. +akai, 9. Jrano +he !olue of the toad edial aydala%anterior preoptic cople# isse#ually diorphic and seasonally !ariable Neurosci. Mett., 22 (;82), pp. *4/G*48

    A.A. Eion et al. =!idence for seasonal plasticity in the onadotropin%releasin horone(n>0) syste of the e$eI Chanes in synaptic inputs onto n>0 neurons =ndocrinoloy, /8(;;7), pp. *2G*4    9. Darnea, F. Notteboh Seasonal recruitent of hippocapal neurons in adult free%ranin black%capped chickadees Proc. Natl. 9cad. Sci. J. S. 9., ; (;;2), pp. *7G**    +.1. Sulders et al. Seasonal !ariation in hippocapal !olue in a food%storin bird, the

 black%capped chickadee A. Neurobiol., *7 (;;4), pp. 4G*4    .+. Sith Seasonal plasticity in the son nuclei of $ild rufous%sided to$hees Drain >es.,7/2 (;;5), pp. 7;G84    9nthony -. +raontin, =liot 9. Dreno$it Seasonal plasticity in the adult brain. +rends in

 Neurosciences, 1olue */, <ssue 5, Aune *, Paes *4G*48

    "+he Principles of Psycholoy", Willia Aaes 8;, Chapter <1, 0abits    Me-ou#, Aoseph =. (**). Synaptic sel! how our brains become who we are. Ne$ Qork,Jnited StatesI 1ikin. p. /7. <SDN %57%/*8%7.    Sha$, Christopher3 :c=achern, Aill, eds. (*). #oward a theory o! neuroplasticity.Mondon, =nlandI Psycholoy Press. <SDN ;78%%825;%*%5.    :ehan ?6>ourke +rain Qour Drain *4 9pril *7    >osen$ei, :ark >. (;;5). "9spects of the search for neural echaniss of eory".

 %nnual &e'iew o! $sychology ,/I G/*. doiI.25@annure!.psych.27... P:<- 85*2/2.    *rain Science $odcast  =pisode Y, "Neuroplasticity"    "Wired Science . 1ideoI :i#ed Feelins". PDS. >etrie!ed * Aune *.    "Shepherd <!ory Fran". >kthoas.y$eb.ua.edu. >etrie!ed * Aune *.

    Colotla, 1ictor 9.3 Dach%y%>ita, Paul (**). "Shepherd <!ory FranI 0is contributions toneuropsycholoy and rehabilitation" (P-F). Cogniti'e, %!!ecti'e *eha'ioral Neuroscience & (*)I 2G28. doiI./748@C9DN.*.*.2.    :auire, =. 9.3 Fracko$iak, >. S.3 Frith, C. -. (;;7). ">ecallin routes around londonI9cti!ation of the riht hippocapus in ta#i dri!ers". #he ournal o! neuroscience the o!!icial

 ournal o! the Society !or Neuroscience +/ (8)I 7/G7. P:<- ;*78422.    Woollett, B.3 :auire, =. 9. (*). "9cHuirin "the Bno$lede" of Mondon6s Mayout-ri!es Structural Drain Chanes". Current *iology &+ (*2)I *;G*2.doiI.5@j.cub.*..8. P:C /*58/45. P:<- **5;4/7. 

. 

  :auire, =. 9.3 adian, -. .3 Aohnsrude, <. S.3 ood, C. -.3 9shburner, A.3Fracko$iak, >. S. A.3 Frith, C. -. (*). "Na!iation%related structural chane in thehippocapi of ta#i dri!ers". $roceedings o! the National %cademy o! Sciences / (8)I2/;8G22/. DibcodeI*PN9S...;7.2/;8:. doiI.7/@pnas.7/;4;7. P:C 8*4/.P:<- 757/8.

urther reading

• Pinaud, >aphael3 +reere, Miisa 9.3 -e Weerd, Peter, eds. (*5).  $lasticity in the 'isual

 system !rom genes to circuits. Ne$ QorkI Spriner. <SDN ;78%%/87%*8;%*.

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• Pinaud, >aphael3 +reere, Miisa 9., eds. (*5). +mmediate early genes in sensory

 processing, cogniti'e per!ormance and neurological disorders. Ne$ QorkI Spriner.<SDN ;78%%/87%//5/%;.

• Deley, Sharon (4 No!eber *2). "Scans of :onks6 Drains Sho$ :editation 9ltersStructure, Functionin". #he 3all Street ournal  (Washinton -.C.). p. D.

• -onohue, Aohn P. (**). "Connectin corte# to achinesI recent ad!ances in braininterfaces" (P-F). Nature Neuroscience .I 84G88. doiI./8@nn;27.P:<- *2/;;*. >etrie!ed February *.

• Flor, 0. (Auly **). "Phanto%lib painI characteristics, causes, and treatent". #he

 (ancet Neurology (=lse!ier) + (/)I 8*G8;. doiI.5@S272%22**(*)72% .

• >aachandran, 1ilayanur S.3 0irstein, Willia (;;8). "+he perception of phantolibs. +he -. ?. 0ebb lecture" (P-F). *rain +&+ (;)I 5/G5/.

doiI.;/@brain@*.;.5/. P:<- ;75*;4*. >etrie!ed / Aanuary *.

• Cohen, Wendy3 0odson, 9nn3 ?60are, 9nne3 Doyle, Aaes3 -urrani, +ariH3 :cCartney,=lspeth3 :attey, :ike3 Naftalin, Mionel3 Watson, Aocelynne (Aune *4). "=ffects ofCoputer%Dased <nter!ention +hrouh 9coustically :odified Speech (Fast ForWord) inSe!ere :i#ed >ecepti!e%=#pressi!e Manuae <pairentI ?utcoes Fro a>andoied Controlled +rial". ournal o! Speech, (anguage, and earing &esearch ,' (/)I 74G7*;. doiI.22@;*%2/88(*4@2;) .

• ister, Sion F. (Aanuary *8). "SC<I Present and Future +herapeutic -e!ices andProstheses". Neurotherapeutics (=lse!ier) . ()I 27G5*.doiI.5@j.nurt.*7..5* . P:C */;874. P:<- 8522;2.

• :ahncke, 0enry W.3 Connor, Donnie D.3 9ppelan, Aed3 9hsanuddin, ?ar N.3 0ardy,Aoseph M.3 Wood, >ichard 9.3 Aoyce, Nicholas :.3 Doniske, +ania et al. (4 9uust *5).":eory enhanceent in healthy older adults usin a brain plasticity%based trainin

 proraI 9 randoied, controlled study". $roceedings o! the National %cademy o!

Sciences o! the 5nited States o! %merica +)( (//)I *4*/G*4*8.doiI.7/@pnas.54;2/. P:C 4*552;. P:<- 5888/8.

• Stein, -onald .3 0offan, Stuart W. (AulyG9uust */). "Concepts of CNS Plasticity

in the Conte#t of Drain -aae and >epair". ournal o! ead #rauma &ehabilitation +' (2)I /7G/2. doiI.;7@;;%*/7%2. P:<- 5****8.

•  Nudo, >andolph A.3 :illiken, arrett W. (;;5). ">eoraniation of :o!eent>epresentations in Priary :otor Corte# Follo$in Focal <scheic <nfarct in 9dultSHuirrel :onkeys". ournal o! Neurophysiology /. (4)I *22G2;. P:<- 87/25.

• Wieloch, +adeus3 Nikolich, Baroly (Aune *5). ":echaniss of neural plasticityfollo$in brain injury". Current )pinion in Neurobiology +* (/)I *48G*52.doiI.5@j.conb.*5.4.. P:<- 57/*24.

1ideos

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• >aachandran. $hantom (imb Syndrome. about consciousness, irror neurons, and phanto lib syndroe

?ther readins

• Chorost, :ichael (*4). &ebuilt how becoming part computer made me more human.

DostonI 0ouhton :ifflin. <SDN %58%/78*;%2.

#xternal lin0s

•  Neuroplasticity at the JS National Mibrary of :edicine :edical Subject 0eadins (:eS0)

•  Neuro :ythsI Separatin Fact and Fiction in Drain%Dased Mearnin by Sara Dernard

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