Aerobic exercise improves cognitive and motor function by inducing neural changes detected using molecular, cellular, and systems level neuroscience techniques. This review unifies the knowledge gained across various neuroscience techniques to provide a comprehensive profile of the neural mechanisms that mediate exercise-induced neuroplasticity. Using a model of exercise-induced neuroplasticity, this review emphasizes the sequence of neural events that accompany exercise, and ultimately promote changes in human performance. This is achieved by differentiating between neuroplasticity induced by acute versus chronic aerobic exercise. Furthermore, this review emphasizes experimental considerations that influence the opportunity to observe exercise-induced neuroplasticity in humans. These include modifiable factors associated with the exercise intervention and nonmodifiable factors such as biological sex, ovarian hormones, genetic variations, and fitness level. To maximize the beneficial effects of exercise in health, disease, and following injury, future research should continue to explore the mechanisms that mediate exercise-induced neuroplasticity. This review identifies some fundamental gaps in knowledge that may serve to guide future research in this area.
AchesonAConoverJCFandlJPDeChiaraTMRussellMThadaniA, and others. 1995. A BDNF autocrine loop in adult sensory neurons prevents cell death. Nature374(6521):450–3.
2.
AmadiUAllmanCJohansen-BergHStaggCJ. 2015. The homeostatic interaction between anodal transcranial direct current stimulation and motor learning in humans is related to GABAA activity. Brain Stimul8(5):898–905.
3.
ArdawiMSRouziAAQariMH. 2012. Physical activity in relation to serum sclerostin, insulin-like growth factor-1, and bone turnover markers in healthy premenopausal women: a cross-sectional and a longitudinal study. J Clin Endocrinol Metab97(10):3691–9.
4.
BaileySPHallEEFolgerSEMillerPC. 2008. Changes in EEG during graded exercise on a recumbent cycle ergometer. J Sports Sci Med7(4):505–11.
5.
BanfiGMarinelliMRoiGSColombiniAPontilloMGiacomettiM, and others. 1994. Growth hormone and insulin-like growth factor I in athletes performing a marathon at 4000 m of altitude. Growth Regul4(2):82–6.
6.
BarhaCKDavisJCFalckRSNagamastsuLSLui-AmbroseT.2017a. Sex differences in exercise efficacy to improve cognition: a systematic review and meta-analysis of randomized controlled trials in older humans. Front Neuroendocrinol46:71–85.
7.
BarhaCKGaleaLANagamastuLSEricksonKILiu-AmbroseT.2017b. Personalising exercise recommendations for brain health: considerations and future directions. Br J Sports Med51(8):636–9.
8.
BegliuominiSCasarosaEPluchinoNLenziECentofantiMFreschiL, and others. 2007. Influence of endogenous and exogenous sex hormones on plasma brain-derived neurotrophic factor. Hum Reprod22(4):995–1002.
9.
BenedictCBrooksSJKullbergJNordenskjoldRBurgosJLe GrevesM, and others. 2013. Association between physical activity and brain health in older adults. Neurobiol Aging34(1):83–90.
10.
BenraissAChmielnickiELernerKRohDGoldmanSA. 2001. Adenoviral brain-derived neurotrophic factor induces both neostriatal and olfactory neuronal recruitment from endogenous progenitor cells in the adult forebrain. J Neurosci21(17):6718–31.
11.
BerchicciMPontifexMBDrolletteESPesceCHillmanCHDi RussoF.2015. From cognitive motor preparation to visual processing: the benefits of childhood fitness to brain health. Neuroscience298:211–9.
12.
BerrettaATzengYCClarksonAN. 2014. Post-stroke recovery: the role of activity-dependent release of brain-derived neurotrophic factor. Expert Rev Neurother14(11):1335–44.
BoraxbekkCJSalamiAWahlinANybergL.2016. Physical activity over a decade modifies age-related decline in perfusion, gray matter volume, and functional connectivity of the posterior default-mode network—a multimodal approach. Neuroimage131:133–41.
15.
BrugniauxJVMarleyCJHodsonDANewKJBaileyDM. 2014. Acute exercise stress reveals cerebrovascular benefits associated with moderate gains in cardiorespiratory fitness. J Cereb Blood Flow Metab34(12):1873–6.
16.
BrunigIPenschuckSBerningerBBensonJFritschyJM. 2001. BDNF reduces miniature inhibitory postsynaptic currents by rapid downregulation of GABA(A) receptor surface expression. Eur J Neurosci13(7):1320–8.
17.
BurdetteJHLaurientiPJEspelandMAMorganATelesfordQVechlekarCD, and others. 2010. Using network science to evaluate exercise-associated brain changes in older adults. Front Aging Neurosci2:23.
18.
BurzynskaAZChaddock-HeymanLVossMWWongCNGotheNPOlsonEA, and others. 2014. Physical activity and cardiorespiratory fitness are beneficial for white matter in low-fit older adults. PLoS One9(9):e107413.
19.
CapponJBraselJAMohanSCooperDM. 1994. Effect of brief exercise on circulating insulin-like growth factor I. J Appl Physiol (1985)76(6):2490–6.
20.
CarsonMJBehringerRRBrinsterRLMcMorrisFA. 1993. Insulin-like growth factor I increases brain growth and central nervous system myelination in transgenic mice. Neuron10(4):729–40.
21.
ChangYKPanCYChenFTTsaiCLHuangCC. 2012. Effect of resistance-exercise training on cognitive function in healthy older adults: a review. J Aging Phys Act20(4):497–517.
22.
ChangYKPesceCChiangYTKuoCYFongDY. 2015. Antecedent acute cycling exercise affects attention control: an ERP study using attention network test. Front Hum Neurosci9:156.
23.
ChapmanSBAslanSSpenceJSDefinaLFKeeblerMWDidehbaniN, and others. 2013. Shorter term aerobic exercise improves brain, cognition, and cardiovascular fitness in aging. Front Aging Neurosci5:75.
24.
ChengACoksayganTTangHKhatriRBalice-GordonRJRaoMS, and others. 2007. Truncated tyrosine kinase B brain-derived neurotrophic factor receptor directs cortical neural stem cells to a glial cell fate by a novel signaling mechanism. J Neurochem100(6):1515–30.
25.
ChirlesTJReiterKWeissLRAlfiniAJNielsonKASmithJC. 2017. Exercise training and functional connectivity changes in mild cognitive impairment and healthy elders. J Alzheimers Dis57(3):845–56.
26.
ChristensenLOJohannsenPSinkjaerTPetersenNPyndtHSNielsenJB. 2000. Cerebral activation during bicycle movements in man. Exp Brain Res135(1):66–72.
27.
ColcombeSKramerAF. 2003. Fitness effects on the cognitive function of older adults: a meta-analytic study. Psychol Sci14(2):125–30.
28.
ColcombeSJEricksonKIRazNWebbAGCohenNJMcAuleyE, and others. 2003. Aerobic fitness reduces brain tissue loss in aging humans. J Gerontol A Biol Sci Med Sci58(2):176–80.
29.
ColcombeSJEricksonKIScalfPEKimJSPrakashRMcAuleyE, and others. 2006. Aerobic exercise training increases brain volume in aging humans. J Gerontol A Biol Sci Med Sci61(11):1166–70.
30.
ColcombeSJKramerAFEricksonKIScalfPMcAuleyECohenNJ, and others. 2004. Cardiovascular fitness, cortical plasticity, and aging. Proc Natl Acad Sci U S A101(9):3316–21.
31.
CosgroveKPMazureCMStaleyJK. 2007. Evolving knowledge of sex differences in brain structure, function and chemistry. Biol Psychiatry62(8):847–55.
32.
CotmanCWBerchtoldNCChristieLA. 2007. Exercise builds brain health: key roles of growth factor cascades and inflammation. Trends Neurosci30(9):464–72.
33.
CurrieJRamsbottomRLudlowHNevillAGilderM.2009. Cardio-respiratory fitness, habitual physical activity and serum brain derived neurotrophic factor (BDNF) in men and women. Neurosci Lett451(2):152–5.
34.
Czarkowska-PaczekBBartlomiejczykIPrzybylskiJ.2006. The serum levels of growth factors: PDGF, TGF-beta and VEGF are increased after strenuous physical exercise. J Physiol Pharmacol57(2):189–97.
35.
DennisAThomasAGRawlingsNBNearJNicholsTEClareS, and others. 2015. An ultra-high field magnetic resonance spectroscopy study of post exercise lactate, glutamate and glutamine change in the human brain. Front Physiol6:351.
36.
DingYHLiJZhouYRafolsJAClarkJCDingY.2006. Cerebral angiogenesis and expression of angiogenic factors in aging rats after exercise. Curr Neurovasc Res3(1):15–23.
37.
DugganCXiaoLWangCYMcTiernanA.2014. Effect of a 12-month exercise intervention on serum biomarkers of angiogenesis in postmenopausal women: a randomized controlled trial. Cancer Epidemiol Biomarkers Prev23(4):648–57.
38.
Duncan-JohnsonCC.1981. P3 Latency: a new metric of information processing. Psychophysiology18:207–15.
39.
EganMFKojimaMCallicottJHGoldbergTEKolachanaBSBertolinoA, and others. 2003. The BDNF val66met polymorphism affects activity-dependent secretion of BDNF and human memory and hippocampal function. Cell112(2):257–69.
EricksonKIPrakashRSVossMWChaddockLHuLMorrisKS, and others. 2009. Aerobic fitness is associated with hippocampal volume in elderly humans. Hippocampus19(10):1030–9.
42.
EricksonKIVossMWPrakashRSBasakCSzaboAChaddockL, and others. 2011. Exercise training increases size of hippocampus and improves memory. Proc Natl Acad Sci U S A108(7):3017–22.
43.
EtnierJLLandersDM. 1998. Motor performance and motor learning as a function of age and fitness. Res Q Exerc Sport69(2):136–46.
44.
FabelKTamBKauferDBaikerASimmonsNKuoCJ, and others. 2003. VEGF is necessary for exercise-induced adult hippocampal neurogenesis. Eur J Neurosci18(10):2803–12.
45.
FarrerLACupplesLAHainesJLHymanBKukullWAMayeuxR, and others. 1997. Effects of age, sex, and ethnicity on the association between apolipoprotein E genotype and Alzheimer disease. A meta-analysis. APOE and Alzheimer Disease Meta Analysis Consortium. JAMA278(16):1349–56.
46.
FerrisLTWilliamsJSShenCL. 2007. The effect of acute exercise on serum brain-derived neurotrophic factor levels and cognitive function. Med Sci Sports Exerc39(4):728–34.
47.
FletcherMALowKABoydRZimmermanBGordonBATanCH, and others. 2016. Comparing aging and fitness effects on brain anatomy. Front Hum Neurosci10:286.
48.
FloelARuscheweyhRKrugerKWillemerCWinterBVolkerK, and others. 2010. Physical activity and memory functions: are neurotrophins and cerebral gray matter volume the missing link?Neuroimage49(3):2756–63.
49.
FloryJDManuckSBFerrellRERyanCMMuldoonMF. 2000. Memory performance and the apolipoprotein E polymorphism in a community sample of middle-aged adults. Am J Med Genet96(6):707–11.
50.
FukuyamaHOuchiYMatsuzakiSNagahamaYYamauchiHOgawaM, and others. 1997. Brain functional activity during gait in normal subjects: a SPECT study. Neurosci Lett228(3):183–6.
51.
GajewskiPDFalkensteinM.2015a. Lifelong physical activity and executive functions in older age assessed by memory based task switching. Neuropsychologia73:195–207.
52.
GajewskiPDFalkensteinM.2015b. Long-term habitual physical activity is associated with lower distractibility in a Stroop interference task in aging: Behavioral and ERP evidence. Brain Cogn98:87–101.
53.
GavinTPRobinsonCBYeagerRCEnglandJANifongLWHicknerRC. 2004. Angiogenic growth factor response to acute systemic exercise in human skeletal muscle. J Appl Physiol (1985)96(1):19–24.
54.
GilesGEBrunyeTTEddyMDMahoneyCRGagnonSATaylorHA, and others. 2014. Acute exercise increases oxygenated and deoxygenated hemoglobin in the prefrontal cortex. Neuroreport25(16):1320–5.
55.
GondohYSensuiHKinomuraSFukudaHFujimotoTMasudM, and others. 2009. Effects of aerobic exercise training on brain structure and psychological well-being in young adults. J Sports Med Phys Fitness49(2):129–35.
56.
GoodmanJMBurrJFBanksLThomasSG. 2016. The acute risks of exercise in apparently healthy adults and relevance for prevention of cardiovascular events. Can J Cardiol32(4):523–32.
57.
GriffinEWMullallySFoleyCWarmingtonSAO’MaraSMKellyAM. 2011. Aerobic exercise improves hippocampal function and increases BDNF in the serum of young adult males. Physiol Behav104(5):934–41.
58.
GungaHCKirschKRockerLBehnCKoralewskiEDavilaEH, and others. 1999. Vascular endothelial growth factor in exercising humans under different environmental conditions. Eur J Appl Physiol Occup Physiol79(6):484–90.
59.
GutmannBMierauAHulsdunkerTHildebrandCPrzyklenkAHollmannW, and others. 2015. Effects of physical exercise on individual resting state EEG alpha peak frequency. Neural Plast2015:717312.
60.
HausmannMTegenthoffMSangerJJanssenFGunturkunOSchwenkreisP.2006. Transcallosal inhibition across the menstrual cycle: a TMS study. Clin Neurophysiol117(1):26–32.
61.
HayesSMHayesJPCaddenMVerfaellieM.2013. A review of cardiorespiratory fitness-related neuroplasticity in the aging brain. Front Aging Neurosci5:31.
62.
HillmanCHBelopolskyAVSnookEMKramerAFMcAuleyE.2004. Physical activity and executive control: implications for increased cognitive health during older adulthood. Res Q Exerc Sport75(2):176–85.
HoltzmanDMPitasREKilbridgeJNathanBMahleyRWBuG, and others. 1995. Low density lipoprotein receptor-related protein mediates apolipoprotein E-dependent neurite outgrowth in a central nervous system-derived neuronal cell line. Proc Natl Acad Sci U S A92:9480–4.
65.
HopkinsMEDavisFCVantieghemMRWhalenPJBucciDJ. 2012. Differential effects of acute and regular physical exercise on cognition and affect. Neuroscience215:59–68.
66.
HsiehJAimoneJBKasparBKKuwabaraTNakashimaKGageFH. 2004. IGF-I instructs multipotent adult neural progenitor cells to become oligodendrocytes. J Cell Biol164(1):111–22.
67.
HuangTLarsenKTRied-LarsenMMollerNCAndersenLB. 2014. The effects of physical activity and exercise on brain-derived neurotrophic factor in healthy humans: a review. Scand J Med Sci Sports24(1):1–10.
68.
HübnerLVoelcker-RehageC.2017. Does physical activity benefit motor performance and learning of upper extremity tasks in older adults? A systematic review. Eur Rev Aging Phys Act14:15.
IeraciAMadaioAIMalleiALeeFSPopoliM.2016. Brain-derived neurotrophic factor Val66Met human polymorphism impairs the beneficial exercise-induced neurobiological changes in mice. Neuropsychopharmacology41(13):3070–9.
71.
InghilleriMConteACurraAFrascaVLorenzanoCBerardelliA.2004. Ovarian hormones and cortical excitability. An rTMS study in humans. Clin Neurophysiol115(5):1063–8.
72.
JahreisGHesseVSchmidtHEScheibeJ.1989. Effect of endurance exercise on somatomedin-C/insulin-like growth factor I concentration in male and female runners. Exp Clin Endocrinol94(1–2):89–96.
73.
JinKZhuYSunYMaoXOXieLGreenbergDA. 2002. Vascular endothelial growth factor (VEGF) stimulates neurogenesis in vitro and in vivo. Proc Natl Acad Sci U S A99(18):11946–50.
74.
JohnsonNFKimCClaseyJLBaileyAGoldBT. 2012. Cardiorespiratory fitness is positively correlated with cerebral white matter integrity in healthy seniors. Neuroimage59(2):1514–23.
75.
JonassonLSNybergLKramerAFLundquistARiklundKBoraxbekkCJ. 2016. Aerobic exercise intervention, cognitive performance, and brain structure: results from the physical influences on brain in aging (PHIBRA) study. Front Aging Neurosci8:336.
76.
JurimaeJTillmannVPurgePJurimaeT.2017. Body composition, maximal aerobic performance and inflammatory biomarkers in endurance-trained athletes. Clin Physiol Funct Imaging37(3):288–92.
77.
KidgellDJPearceAJ. 2011. What has transcranial magnetic stimulation taught us about neural adaptations to strength training? A brief review. J Strength Cond Res25(11):3208–17.
78.
KleemeyerMMKuhnSPrindleJBodammerNCBrechtelLGartheA, and others. 2016. Changes in fitness are associated with changes in hippocampal microstructure and hippocampal volume among older adults. Neuroimage131:155–61.
79.
KleimJAChanSPringleESchallertKProcaccioVJimenezR, and others. 2006. BDNF val66met polymorphism is associated with modified experience-dependent plasticity in human motor cortex. Nat Neurosci9(6):735–7.
80.
KlimeschW.2012. Alpha-band oscillations, attention, and controlled access to stored information. Trends Cogn Sci16(12):606–17.
81.
KoistinenHKoistinenRSeleniusLYlikorkalaQSeppalaM.1996. Effect of marathon run on serum IGF-I and IGF-binding protein 1 and 3 levels. J Appl Physiol (1985)80(3):760–4.
82.
KowianskiPLietzauGCzubaEWaskowMSteligaAMorysJ.2017. BDNF: a key factor with multipotent impact on brain signaling and synaptic plasticity. Cell Mol Neurobiol38(3):579–93.
83.
KraemerRRDurandRJAcevedoEOJohnsonLGKraemerGRHebertEP, and others. 2004. Rigorous running increases growth hormone and insulin-like growth factor-I without altering ghrelin. Exp Biol Med (Maywood)229(3):240–6.
84.
KrausRMStallingsHW3rdYeagerRCGavinTP. 2004. Circulating plasma VEGF response to exercise in sedentary and endurance-trained men. J Appl Physiol (1985)96(4):1445–50.
85.
KuoMFPaulusWNitscheMA. 2006. Sex differences in cortical neuroplasticity in humans. Neuroreport17(16):1703–7.
86.
LangeKHIsakssonFJuulARasmussenMHBulowJKjaerM.2000. Growth hormone enhances effects of endurance training on oxidative muscle metabolism in elderly women. Am J Physiol Endocrinol Metab279(5):E989–96.
87.
LeckieRLOberlinLEVossMWPrakashRSSzabo-ReedAChaddock-HeymanL, and others. 2014. BDNF mediates improvements in executive function following a 1-year exercise intervention. Front Hum Neurosci8:985.
88.
LeffertsWKBabcockMCTissMJIvesSJWhiteCNBrutsaertTD, and others. 2016. Effect of hypoxia on cerebrovascular and cognitive function during moderate intensity exercise. Physiol Behav165:108–18.
89.
LemosJRJrAlvesCRde SouzaSBMarsigliaJDSilvaMSPereiraAC, and others. 2016. Peripheral vascular reactivity and serum BDNF responses to aerobic training are impaired by the BDNF Val66Met polymorphism. Physiol Genomics48(2):116–23.
90.
LiLMenWWChangYKFanMXJiLWeiGX. 2014. Acute aerobic exercise increases cortical activity during working memory: a functional MRI study in female college students. PLoS One9(6):e99222.
91.
LinCYHungSYChenHTTsouHKFongYCWangSW, and others. 2014. Brain-derived neurotrophic factor increases vascular endothelial growth factor expression and enhances angiogenesis in human chondrosarcoma cells. Biochem Pharmacol91(4):522–33.
92.
ListaISorrentinoG.2010. Biological mechanisms of physical activity in preventing cognitive decline. Cell Mol Neurobiol30(4):493–503.
93.
Lopez-LopezCLeRoithDTorres-AlemanI.2004. Insulin-like growth factor I is required for vessel remodeling in the adult brain. Proc Natl Acad Sci U S A101(26):9833–8.
94.
LouissaintAJrRaoSLeventhalCGoldmanSA. 2002. Coordinated interaction of neurogenesis and angiogenesis in the adult songbird brain. Neuron34(6):945–60.
95.
LucasSJAinsliePNMurrellCJThomasKNFranzEACotterJD. 2012. Effect of age on exercise-induced alterations in cognitive executive function: relationship to cerebral perfusion. Exp Gerontol47(8):541–51.
MaassADuzelSBrigadskiTGoerkeMBeckeASobierayU, and others. 2016. Relationships of peripheral IGF-1, VEGF and BDNF levels to exercise-related changes in memory, hippocampal perfusion and volumes in older adults. Neuroimage131:142–54.
98.
MaassADuzelSGoerkeMBeckeASobierayUNeumannK, and others. 2015. Vascular hippocampal plasticity after aerobic exercise in older adults. Mol Psychiatry20(5):585–93.
99.
MacIntoshBJCraneDESageMDRajabASDonahueMJMcIlroyWE, and others. 2014. Impact of a single bout of aerobic exercise on regional brain perfusion and activation responses in healthy young adults. PLoS One9(1):e85163.
100.
MaddockRJCasazzaGABuonocoreMHTanaseC.2011. Vigorous exercise increases brain lactate and Glx (glutamate + glutamine): a dynamic 1H-MRS study. Neuroimage57(4):1324–30.
101.
MaddockRJCasazzaGAFernandezDHMaddockMI. 2016. Acute modulation of cortical glutamate and GABA content by physical activity. J Neurosci36(8):2449–57.
102.
MakMKWong-YuISShenXChungCL. 2017. Long-term effects of exercise and physical therapy in people with Parkinson disease. Nat Rev Neurol13(11): 689–703.
103.
MandyamCDWeeSEischAJRichardsonHNKoobGF. 2007. Methamphetamine self-administration and voluntary exercise have opposing effects on medial prefrontal cortex gliogenesis. J Neurosci27(42):11442–50.
104.
ManettaJBrunJFMaimounLFedouCPrefautCMercierJ.2003. The effects of intensive training on insulin-like growth factor I (IGF-I) and IGF binding proteins 1 and 3 in competitive cyclists: relationships with glucose disposal. J Sports Sci21(3):147–54.
105.
MangCSBrownKENevaJLSnowNJCampbellKLBoydLA. 2016. Promoting motor cortical plasticity with acute aerobic exercise: a role for cerebellar circuits. Neural Plast2016:6797928.
106.
MangCSCampbellKLRossCJBoydLA. 2013. Promoting neuroplasticity for motor rehabilitation after stroke: considering the effects of aerobic exercise and genetic variation on brain-derived neurotrophic factor. Phys Ther93(12):1707–16.
107.
MangCSMcEwenLMMacIsaacJLSnowNJCampbellKLKoborMS, and others. 2017. Exploring genetic influences of underlying acute aerobic exercise effects on motor learning. Sci Rep7(1):12123.
108.
McEwenBSAlvesSEBullochKWeilandNG. 1997. Ovarian steroids and the brain: implications for cognition and aging. Neurology48(5Suppl 7):S8–15.
109.
McGregorKMNoceraJRSudhyadhomAPattenCManiniTMKleimJA, and others. 2013. Effects of aerobic fitness on aging-related changes of interhemispheric inhibition and motor performance. Front Aging Neurosci5:66.
110.
McGregorKMZlatarZKleimESudhyadhomABauerAPhanS, and others. 2011. Physical activity and neural correlates of aging: a combined TMS/fMRI study. Behav Brain Res222(1):158–68.
111.
McHughenSARodriguezPFKleimJAKleimEDMarchal CrespoLProcaccioV, and others. 2017. BDNF val66met polymorphism influences motor system function in the human brain. Cereb Cortex20(5):1254–62.
112.
MiddletonLKirklandSRockwoodK.2008. Prevention of CIND by physical activity: different impact on VCI-ND compared with MCI. J Neurol Sci269(1–2):80–4.
113.
MonteggiaLMBarrotMPowellCMBertonOGalanisVGemelliT, and others. 2004. Essential role of brain-derived neurotrophic factor in adult hippocampal function. Proc Natl Acad Sci U S A101(29):10827–32.
MoraesHFerreiraCDeslandesACagyMPompeuFRibeiroP, and others. 2007. Beta and alpha electroencephalographic activity changes after acute exercise. Arq Neuropsiquiatr65(3a):637–41.
116.
NandaBBaldeJManjunathaS.2013. The acute effects of a single bout of moderate-intensity aerobic exercise on cognitive functions in healthy adult males. J Clin Diagn Res7(9):1883–5.
117.
Nieto-EstevezVDefteraliCVicario-AbejonC.2016. IGF-I: a key growth factor that regulates neurogenesis and synaptogenesis from embryonic to adult stages of the brain. Front Neurosci10:52.
118.
NindlBCScofieldDEStrohbachCACentiAJEvansRKYanovichR, and others. 2012. IGF-I, IGFBPs, and inflammatory cytokine responses during gender-integrated Israeli Army basic combat training. J Strength Cond Res26(Suppl 2):S73–81.
119.
NofujiYSuwaMSasakiHIchimiyaANishichiRKumagaiS.2012. Different circulating brain-derived neurotrophic factor responses to acute exercise between physically active and sedentary subjects. J Sports Sci Med11(1):83–8.
120.
OberlinLEVerstynenTDBurzynskaAZVossMWPrakashRSChaddock-HeymanL, and others. 2016. White matter microstructure mediates the relationship between cardiorespiratory fitness and spatial working memory in older adults. Neuroimage131:91–101.
121.
Ozyemisci-TaskiranOGunendiZBolukbasiNBeyazovaM.2008. The effect of a single session submaximal aerobic exercise on premotor fraction of reaction time: an electromyographic study. Clin Biomech (Bristol, Avon)23(2):231–5.
122.
Pearson-FuhrhopKMCramerSC. 2010. Genetic influences on neural plasticity. PM R2(12Suppl 2):S227–40.
123.
PenceaVBingamanKDWiegandSJLuskinMB. 2001. Infusion of brain-derived neurotrophic factor into the lateral ventricle of the adult rat leads to new neurons in the parenchyma of the striatum, septum, thalamus, and hypothalamus. J Neurosci21(17):6706–17.
124.
Perez-MartinMCifuentesMGrondonaJMLopez-AvalosMDGomez-PinedoUGarcia-VerdugoJM, and others. 2010. IGF-I stimulates neurogenesis in the hypothalamus of adult rats. Eur J Neurosci31(9):1533–48.
125.
PetruzzelloSJHallEEEkkekakisP.2001. Regional brain activation as a biological marker of affective responsivity to acute exercise: influence of fitness. Psychophysiology38(1):99–106.
126.
PolichJHeineMRD. 1996. P3 topography and modality effects from a single-stimulus paradigm. Psychophysiology103:493–8.
127.
PooMM.2001. Neurotrophins as synaptic modulators. Nat Rev Neurosci2(1):24–32.
128.
PrakashRSVossMWEricksonKILewisJMChaddockLMalkowskiE, and others. 2011. Cardiorespiratory fitness and attentional control in the aging brain. Front Hum Neurosci4:229.
129.
RaichlenDABharadwajPKFitzhughMCHawsKATorreGATrouardTP, and others. 2016. Differences in resting state functional connectivity between young adult endurance athletes and healthy controls. Front Hum Neurosci10:610.
130.
RajabASCraneDEMiddletonLERobertsonADHampsonMMacIntoshBJ. 2014. A single session of exercise increases connectivity in sensorimotor-related brain networks: a resting-state fMRI study in young healthy adults. Front Hum Neurosci8:625.
131.
RiddingMCZiemannU.2010. Determinants of the induction of cortical plasticity by non-invasive brain stimulation in healthy subjects. J Physiol588(Pt 13):2291–304.
132.
RoigMSkriverKLundbye-JensenJKiensBNielsenJB. 2012. A single bout of exercise improves motor memory. PLoS One7(9):e44594.
133.
Rojas VegaSHollmannWVera WahrmannBStrüderHK. 2012. pH buffering does not influence BDNF responses to exercise. Int J Sports Med33:8–12.
134.
RosanoCVenkatramanVKGuralnikJNewmanABGlynnNWLaunerL, and others. 2010. Psychomotor speed and functional brain MRI 2 years after completing a physical activity treatment. J Gerontol A Biol Sci Med Sci65(6):639–47.
135.
RosendalLLangbergHFlyvbjergAFrystykJOrskovHKjaerM.2002. Physical capacity influences the response of insulin-like growth factor and its binding proteins to training. J Appl Physiol (1985)93(5):1669–75.
136.
RossiterHEDavisEMClarkEVBoudriasMHWardNS. 2014. Beta oscillations reflect changes in motor cortex inhibition in healthy ageing. Neuroimage91:360–5.
137.
RudmanDMattsonDE. 1994. Serum insulin-like growth factor I in healthy older men in relation to physical activity. J Am Geriatr Soc42(1):71–6.
138.
RuscheweyhRWillemerCKrugerKDuningTWarneckeTSommerJ, and others. 2011. Physical activity and memory functions: an interventional study. Neurobiol Aging32(7):1304–19.
139.
SchifferTSchulteSHollmannWBlochWStruderHK. 2009. Effects of strength and endurance training on brain-derived neurotrophic factor and insulin-like growth factor 1 in humans. Horm Metab Res41(3):250–4.
140.
Schmidt-KassowMSchadleSOtterbeinSThielCDoehringALotschJ, and others. 2012. Kinetics of serum brain-derived neurotrophic factor following low-intensity versus high-intensity exercise in men and women. Neuroreport23(15):889–93.
141.
SchmoleskyMTWebbDLHansenRA. 2013. The effects of aerobic exercise intensity and duration on levels of brain-derived neurotrophic factor in healthy men. J Sports Sci Med12(3):502–11.
142.
SchneiderSAskewCDDiehlJMierauAKleinertJAbelT, and others. 2009. EEG activity and mood in health orientated runners after different exercise intensities. Physiol Behav96(4–5):709–16.
143.
SchobersbergerWHobisch-HagenPFriesDWiedermannFRieder-ScharingerJVilligerB, and others. 2000. Increase in immune activation, vascular endothelial growth factor and erythropoietin after an ultramarathon run at moderate altitude. Immunobiology201(5):611–20.
144.
SchwarzAJBraselJAHintzRLMohanSCooperDM. 1996. Acute effect of brief low- and high-intensity exercise on circulating insulin-like growth factor (IGF) I, II, and IGF-binding protein-3 and its proteolysis in young healthy men. J Clin Endocrinol Metab81(10):3492–7.
145.
SeifertTBrassardPWissenbergMRasmussenPNordbyPStallknechtB, and others. 2010. Endurance training enhances BDNF release from the human brain. Am J Physiol Regul Integr Comp Physiol298(2):R372–7.
146.
SextonCEBettsJFDemnitzNDawesHEbmeierKPJohansen-BergH.2016. A systematic review of MRI studies examining the relationship between physical fitness and activity and the white matter of the ageing brain. Neuroimage131:81–90.
147.
ShimadaHIshiiKIshiwataKOdaKSuzukawaMMakizakoH, and others. 2013. Gait adaptability and brain activity during unaccustomed treadmill walking in healthy elderly females. Gait Posture38(2):203–8.
148.
SinghAMDuncanRENevaJLStainesWR. 2014. Aerobic exercise modulates intracortical inhibition and facilitation in a nonexercised upper limb muscle. BMC Sports Sci Med Rehabil6:23.
149.
SkriverKRoigMLundbye-JensenJPingelJHelgeJWKiensB, and others. 2014. Acute exercise improves motor memory: exploring potential biomarkers. Neurobiol Learn Mem116:46–58.
150.
SmithAEGoldsworthyMRGarsideTWoodFMRiddingMC. 2014. The influence of a single bout of aerobic exercise on short-interval intracortical excitability. Exp Brain Res232(6):1875–82.
151.
SmithJCPaulsonESCookDBVerberMDTianQ.2010. Detecting changes in human cerebral blood flow after acute exercise using arterial spin labeling: implications for fMRI. J Neurosci Methods191(2):258–62.
152.
SmithMJAdamsLFSchmidtPJRubinowDRWassermannEM. 2002. Effects of ovarian hormones on human cortical excitability. Ann Neurol51(5):599–603.
153.
SmithMJKeelJCGreenbergBDAdamsLFSchmidtPJRubinowDA, and others. 1999. Menstrual cycle effects on cortical excitability. Neurology53(9):2069–72.
154.
SnowNJMangCSRoigMMcDonnellMNCampbellKLBoydLA. 2016. The effect of an acute bout of moderate-intensity aerobic exercise on motor learning of a continuous tracking task. PLoS One11(2):e0150039.
155.
SohrabjiFLewisDK. 2006. Estrogen-BDNF interactions: implications for neurodegenerative diseases. Front Neuroendocrinol27(4):404–14.
156.
SonntagWEDeakFAshpoleNTothPCsiszarAFreemanW, and others. 2013. Insulin-like growth factor-1 in CNS and cerebrovascular aging. Front Aging Neurosci5:27.
157.
StattonMAEncarnacionMCelnikPBastianAJ. 2015. A single bout of moderate aerobic exercise improves motor skill acquisition. PLoS One10(10):e0141393.
158.
StavrinosELCoxonJP. 2017. High-intensity interval exercise promotes motor cortex disinhibition and early motor skill consolidation. J Cogn Neurosci29(4):593–604.
159.
StillmanCMCohenJLehmanMEEricksonKI. 2016. Mediators of physical activity on neurocognitive function: a review at multiple levels of analysis. Front Hum Neurosci10:626.
160.
SvatkovaAMandlRCScheeweTWCahnWKahnRSHulshoff PolHE. 2015. Physical exercise keeps the brain connected: biking increases white matter integrity in patients with schizophrenia and healthy controls. Schizophr Bull41(4):869–78.
161.
SzuhanyKLBugattiMOttoMW. 2015. A meta-analytic review of the effects of exercise on brain-derived neurotrophic factor. J Psychiatr Res60:56–64.
162.
TashiroMItohMFujimotoTFujiwaraTOtaHKubotaK, and others. 2001. 18F-FDG PET mapping of regional brain activity in runners. J Sports Med Phys Fitness41(1):11–7.
163.
ThomasAGDennisARawlingsNBStaggCJMatthewsLMorrisM, and others. 2016a. Multi-modal characterization of rapid anterior hippocampal volume increase associated with aerobic exercise. Neuroimage131:162–70.
164.
ThomasRBeckMMLindRRKorsgaard JohnesnLGeertsenSSChristiansenL, and others. 2016b. Acute exercise and motor memory consolidation: the role of exercise timing. Neural Plast2016:6205452.
165.
ThomasRFlindtgaardMSkriverKGeertsenSSChristiansenLKorsgaard JohnesnL, and others. 2016c. Acute exercise and motor memory consolidation: does exercise type play a role?Scand J Med Sci Sports27(11):1523–32.
166.
ThomasRJohnsenLKGeertsenSSChristiansenLRitzCRoigM, and others. 2016d. Acute exercise and motor memory consolidation: the role of exercise intensity. PLoS One11(7):e0159589.
167.
ThomasBPYezhuvathUSTsengBYLiuPLevineBDZhangR, and others. 2013. Life-long aerobic exercise preserved baseline cerebral blood flow but reduced vascular reactivity to CO2. J Magn Reson Imaging38(5):1177–83.
168.
TianQEricksonKISimonsickEMAizensteinHJGlynnNWBoudreauRM, and others. 2014a. Physical activity predicts microstructural integrity in memory-related networks in very old adults. J Gerontol A Biol Sci Med Sci69(10):1284–90.
169.
TianQSimonsickEMEricksonKIAizensteinHJGlynnNWBoudreauRM, and others. 2014b. Cardiorespiratory fitness and brain diffusion tensor imaging in adults over 80 years of age. Brain Res1588:63–72.
170.
Torres-AlemanINaftolinFRobbinsRJ. 1990. Trophic effects of basic fibroblast growth factor on fetal rat hypothalamic cells: interactions with insulin-like growth factor I. Brain Res Dev Brain Res52(1–2):253–7.
171.
TrejoJLCarroETorres-AlemanI.2001. Circulating insulin-like growth factor I mediates exercise-induced increases in the number of new neurons in the adult hippocampus. J Neurosci21(5):1628–34.
172.
TrejoJLPirizJLlorens-MartinMVFernandezAMBolosMLeRoithD, and others. 2007. Central actions of liver-derived insulin-like growth factor I underlying its pro-cognitive effects. Mol Psychiatry12(12):1118–28.
173.
TsaiCLPanCYChenFCWangCHChouFY. 2016. Effects of acute aerobic exercise on a task-switching protocol and brain-derived neurotrophic factor concentrations in young adults with different levels of cardiorespiratory fitness. Exp Physiol101(7):836–50.
174.
TsengBYUhJRossettiHCCullumCMDiaz-ArrastiaRFLevineBD, and others. 2013. Masters athletes exhibit larger regional brain volume and better cognitive performance than sedentary older adults. J Magn Reson Imaging38(5):1169–76.
175.
VaaraJPKalliomaaRHynninenPKyrolainenH.2015. Physical fitness and hormonal profile during an 11-week paratroop training period. J Strength Cond Res29(Suppl 11):S163–7.
176.
van PraagH. 2008. Neurogenesis and exercise: past and future directions. Neuromol Med10(2):128–40.
177.
VerkooijenSStevelinkRAbramovicLVinkersCHOphoffRAKahnRS, and others. 2017. The association of sleep and physical activity with integrity of white matter microstructure in bipolar disorder patients and healthy controls. Psychiatry Res262:71–80.
178.
VerstynenTDLynchBMillerDLVossMWPrakashRSChaddockL, and others. 2012. Caudate nucleus volume mediates the link between cardiorespiratory fitness and cognitive flexibility in older adults. J Aging Res2012:939285.
179.
Voelcker-RehageCGoddeBStaudingerUM. 2011. Cardiovascular and coordination training differentially improve cognitive performance and neural processing in older adults. Front Hum Neurosci5:26.
180.
VoskuilDWBuenodeMesquitaHBKaaksRvan NoordPARinaldiSRiboliE, and others. 2001. Determinants of circulating insulin-like growth factor (IGF)-I and IGF binding proteins 1–3 in premenopausal women: physical activity and anthropometry (Netherlands). Cancer Causes Control12(10):951–8.
181.
VossMWEricksonKIPrakashRSChaddockLKimJSAlvesH, and others. 2013a. Neurobiological markers of exercise-related brain plasticity in older adults. Brain Behav Immun28:90–9.
182.
VossMWEricksonKIPrakashRSChaddockLMalkowskiEAlvesH, and others. 2010. Functional connectivity: a source of variance in the association between cardiorespiratory fitness and cognition?Neuropsychologia48(5):1394–406.
183.
VossMWHeoSPrakashRSEricksonKIAlvesHChaddockL, and others. 2013b. The influence of aerobic fitness on cerebral white matter integrity and cognitive function in older adults: results of a one-year exercise intervention. Hum Brain Mapp34(11):2972–85.
184.
VossMWNagamatsuLSLiu-AmbroseTKramerAF. 2011. Exercise, brain, and cognition across the life span. J Appl Physiol (1985)111(5):1505–13.
185.
WagnerGHerbslebMde la CruzFSchumannAKohlerSPutaC, and others. 2017. Changes in fMRI activation in anterior hippocampus and motor cortex during memory retrieval after an intense exercise intervention. Biol Psychol124:65–78.
186.
WahlPZinnerCAchtzehnSBehringerMBlochWMesterJ.2011. Effects of acid-base balance and high or low intensity exercise on VEGF and bFGF. Eur J Appl Physiol111(7):1405–13.
187.
WangJD’AmatoABambroughJSwartzAMMillerNE. 2016. A positive association between active lifestyle and hemispheric lateralization for motor control and learning in older adults. Behav Brain Res314:38–44.
188.
WengTBPierceGLDarlingWGVossMW. 2015. Differential effects of acute exercise on distinct aspects of executive function. Med Sci Sports Exerc47(7):1460–9.
189.
WilliamsonJWMcCollRMathewsDGinsburgMMitchellJH. 1999. Activation of the insular cortex is affected by the intensity of exercise. J Appl Physiol (1985)87(3):1213–9.
190.
WongCNChaddock-HeymanLVossMWBurzynskaAZBasakCEricksonKI, and others. 2015. Brain activation during dual-task processing is associated with cardiorespiratory fitness and performance in older adults. Front Aging Neurosci7:154.
191.
WoodKNNikolovRShoemakerJK. 2016. Impact of long-term endurance training vs. guideline-based physical activity on brain structure in healthy aging. Front Aging Neurosci8:155.
192.
XingYYangSDDongFWangMMFengYSZhangF.2018. The beneficial role of early exercise training following stroke and possible mechanisms. Life Sci198:32–37.
193.
YamaguchiTFujiwaraTLiuWLiuM.2012. Effects of pedaling exercise on the intracortical inhibition of cortical leg area. Exp Brain Res218(3):401–6.
194.
YanagisawaHDanITsuzukiDKatoMOkamotoMKyutokuY, and others. 2010. Acute moderate exercise elicits increased dorsolateral prefrontal activation and improves cognitive performance with Stroop test. Neuroimage50(4):1702–10.
195.
YePCarsonJD’ErcoleAJ. 1995. In vivo actions of insulin-like growth factor-I (IGF-I) on brain myelination: studies of IGF-I and IGF binding protein-1 (IGFBP-1) transgenic mice. J Neurosci15(11):7344–56.
196.
YoungstedtSDDishmanRKCuretonKJPeacockLJ. 1993. Does body temperature mediate anxiolytic effects of acute exercise?J Appl Physiol (1985)74(2):825–31.
197.
ZiemannUReisJSchwenkreisPRosanovaMStrafellaABadawyR, and others. 2014. TMS and drugs revisited 2014. Clin Neurophysiol126(10):1847–68.
198.
ZigovaTPenceaVWiegandSJLuskinMB. 1998. Intraventricular administration of BDNF increases the number of newly generated neurons in the adult olfactory bulb. Mol Cell Neurosci11(4):234–45.
199.
ZimmermanBSuttonBPLowKAFletcherMATanCHSchneider-GarcesN, and others. 2014. Cardiorespiratory fitness mediates the effects of aging on cerebral blood flow. Front Aging Neurosci6:59.
200.
ZoladzJAPilcAMajerczakJGrandysMZapart-BukowskaJDudaK.2008. Endurance training increases plasma brain-derived neurotrophic factor concentration in young healthy men. J Physiol Pharmacol59(Suppl 7):119–32.
