Journal Mobile Options
Table of Contents
Vol. 32, No. 3, 2010
Issue release date: August 2010
Section title: Original Paper
Editor's Choice -- Free Access
Dev Neurosci 2010;32:249–256
(DOI:10.1159/000316648)

Basal Ganglia Volume Is Associated with Aerobic Fitness in Preadolescent Children

Chaddock L.a · Erickson K.I.c · Prakash R.S.d · VanPatter M.a · Voss M.W.a · Pontifex M.B.b · Raine L.B.b · Hillman C.H.b · Kramer A.F.a
aDepartment of Psychology, Beckman Institute for Advanced Science and Technology, and bDepartment of Kinesiology and Community Health, University of Illinois at Urbana-Champaign, Urbana, Ill., cDepartment of Psychology, University of Pittsburgh, Pittsburgh, Pa., and dDepartment of Psychology, Ohio State University, Columbus, Ohio, USA
email Corresponding Author

Abstract

The present investigation is the first to explore the association between childhood aerobic fitness and basal ganglia structure and function. Rodent research has revealed that exercise influences the striatum by increasing dopamine signaling and angiogenesis. In children, higher aerobic fitness levels are associated with greater hippocampal volumes, superior performance on tasks of attentional and interference control, and elevated event-related brain potential indices of executive function. The present study used magnetic resonance imaging to investigate if higher-fit and lower-fit 9- and 10-year-old children exhibited differential volumes of other subcortical brain regions, specifically the basal ganglia involved in attentional control. The relationship between aerobic fitness, dorsal and ventral striatum volumes and performance on an attention and inhibition Eriksen flanker task was also examined. The results indicated that higher-fit children showed superior flanker task performance compared to lower-fit children. Higher-fit children also showed greater volumes of the dorsal striatum, and dorsal striatum volume was negatively associated with behavioral interference. The results support the claim that the dorsal striatum is involved in cognitive control and response resolution and that these cognitive processes vary as a function of aerobic fitness. No relationship was found between aerobic fitness, the volume of the ventral striatum and flanker performance. The findings suggest that increased childhood aerobic fitness is associated with greater dorsal striatal volumes and that this is related to enhanced cognitive control. Because children are becoming increasingly overweight, unhealthy and unfit, understanding the neurocognitive benefits of an active lifestyle during childhood has important public health and educational implications.

© 2010 S. Karger AG, Basel


  

Key Words

  • Brain
  • Development
  • Exercise
  • MRI
  • Physical activity
  • Neurocognition
  • Neuroimaging
  • Striatum

References

  1. Aguiar AS, Speck AE, Prediger RD, Kapczinski F, Pinho RA (2008): Downhill training upregulates mice hippocampal and striatal brain-derived neurotrophic factor levels. J Neural Transm 115:1251–1255.
  2. American College of Sports Medicine (2006): ACSM’s Guidelines for Exercise Testing and Prescription, ed 7. New York, Lippincott Williams & Wilkins, p 366.
  3. Aron AR, Poldrack RA, Wise SP (2009): Cognition: basal ganglia role; in Squire LR (ed.): Encyclopedia of Neuroscience, vol 2, pp 1069–1077.

    External Resources

  4. Baker JL, Olsen LW, Sorensen TIA (2007): Childhood body-mass index and risk of coronary heart disease in adulthood. N Engl J Med 357:2329–2337.
  5. Bar-Or O (1983): Pediatric sports medicine for the practitioner: From physiologic principles to clinical applications. New York, Springer, p 376.
  6. Birnbaum AS, Lytle LA, Murray DM, Story M, Perry CL, Boutelle KN (2002): Survey development for assessing correlates of young adolescents’ eating. Am J Health Behav 26:284–295.
  7. Buck SM, Hillman CH, Castelli DM (2008): The relation of aerobic fitness to Stroop task performance in preadolescent children. Med Sci Sports Exerc 40:166–172.
  8. Casey BJ, Getz S, Galvan A (2008): The adolescent brain. Dev Rev 28:62–77.
  9. Casey BJ, Thomas KM, Welsh TF, Badgaiyan RD, Eccard CH, Jennings JR, Crone EA (2000): Dissociation of response conflict, attentional selection, and expectancy with functional magnetic resonance imaging. Proc Natl Acad Sci 97:8728–8733.
  10. Casey BJ, Trainor RJ, Orendi JL, Schubert AB, Nystrom LE, Giedd JN, Castellanos FX, Haxby JV, Noll DC, Cohen JD, Forman SD, Dahl RE, Rapoport JL (1997): A developmental functional MRI study of prefrontal activation during performance of a go-no-go task. J Cog Neurosci 9:835–847.
  11. Castelli DM, Hillman CH, Buck SM, Erwin HE (2007): Physical fitness and academic achievement in third- and fifth-grade students. J Sport Exerc Psychol 29:239–252.
  12. Chaddock L, Erickson KI, Prakash RS, Kim JS, Voss MW, VanPatter M, Pontifex MB, Raine LB, Konkel A, Hillman CH, Cohen NJ, Kramer AF (2010): A neuroimaging investigation of the association between aerobic fitness, hippocampal volume and memory performance in preadolescent children. Devel- opment & Aging. Abstr Cogn Neurosci Soc B82-76.
  13. Chaddock L, Hillman CH, Buck SM, Cohen NJ (in press): Aerobic fitness and executive control of relational memory in preadolescent children: MSSE.
  14. Chomitz VR, Slining MM, McGowan RJ, Mitchell SE, Dawson GF, Hacker KA (2009): Is there a relationship between physical fitness and academic achievement? Positive results from public school children in the northeastern United States. J Sch Health 79:30–37.
  15. Colcombe S, Kramer AF (2003): Fitness effects on the cognitive function of older adults: A meta-analytic study. Psychol Sci 14:125–130.
  16. Colcombe SJ, Kramer AF, Erickson KI, Scalf P, McAuley E, Cohen NJ, Webb A, Jerome GJ, Marquez DX, Elavsky S (2004): Cardiovascular fitness, cortical plasticity, and aging. PNAS 101:3316–3321.
  17. Cotman CW, Berchtold NC (2002): Exercise: a behavioral intervention to enhance brain health and plasticity. Trends Neurosci 25:295–301.
  18. Di Martino A, Scheres A, Marguiles DS, Kelly AM, Uddin LQ, Shehzad Z, Biswal B, Walters JR, Castellanos FX, Milham MP (2008): Functional connectivity of human striatum: a resting state fMRI study. Cereb Cortex 18:2735–2747.
  19. Ding YH, Luan XD, Li J, Rafols JA, Guthinkonda M, Diaz FG, Ding Y (2004): Exercise-induced overexpression of angiogenic factors and reduction of ischemia/reperfusion injury in stroke. Curr Neurovasc Res 1:411–420.
  20. Draganski B, Kherif F, Kloppel S, Cook PA, Alexander DC, Parker GJ, Deichmann R, Ashburner J, Frackowiak RS (2008): Evidence for segregated and integrative connectivity patterns in the human basal ganglia. J Neurosci 28:7143–7152.
  21. DuPaul GJ, Power TJ, Anastopoulos A, Reid R (1998): ADHD Rating Scale – IV: Checklists, Norms, and Clinical Interpretation. New York, Guilford Press.
  22. Erickson KI, Boot WR, Basak C, Neider MR, Prakash RS, Voss MW, Graybiel AM, Simons DJ, Fabiani M, Gratton G, Kramer AF (in press): Striatal volume predicts level of video game skill acquisition.
  23. Erickson KI, Prakash RS, Voss MW, Chaddock L, Hu L, Morris KS, White SM, Wojcicki TR, McAuley E, Kramer AF (2009): Aerobic fitness is associated with hippocampal volume in elderly humans. Hippocampus 19:1030–1039.
  24. Eriksen BA, Eriksen CW (1974): Effects of noise letters on the identification of a target letter in a nonsearch task. Percept Psychophys 16:143–149.
  25. Freedson PS, Goodman TL (1993): Measurement of oxygen consumption; in Rowland TW (ed): Pediatric Laboratory Exercise Testing: Clinical Guidelines. Human Kinetics, Champaign, pp 91–113.
  26. Graybiel AM (2005): The basal ganglia: learning new tricks and loving it. Curr Opin Neurobiol 15:638–644.
  27. Graybiel AM (2008): Habits, rituals and the evaluative brain. Annu Rev Neurosci 31:359–387.
  28. Hillman CH, Buck SM, Themanson JR, Pontifex MB, Castelli DM (2009): Aerobic fitness and cognitive development: event-related brain potential and task performance of executive control in preadolescent children. Dev Psychol 45:114–129.
  29. Hillman CH, Castelli DM, Buck SM (2005): Aerobic fitness and neurocognitive function in healthy preadolescent children. Med Sci Sports Exerc 37:1967–1974.
  30. Hillman CH, Erickson KI, Kramer AF (2008): Be smart, exercise your heart: exercise effects on brain and cognition. Nature Rev Neurosci 9:58–65.
  31. Kaufman AS, Kaufman NL (1990): Kaufman Brief Intelligence Test. Circle Pines, AGS.
  32. Kramer AF, Hahn S, Cohen N, Banich M, McAuley E, Harrison C, Chason J, Vakil E, Bardell L, Boileau RA, Colcombe A (1999): Aging, fitness, and neurocognitive function. Nature 400:418–419.
  33. Kramer A, Humphrey D, Larish J, Logan G, Strayer D (1994): Aging and inhibition: beyond a unitary view of inhibitory processing in attention. Psychol Aging 9:491–512.
  34. Li J, Ding YH, Rafols JA, Lai Q, McAllister JP, Ding Y (2005): Increased astrocyte proliferation in rats after running exercise. Neurosci Lett 386:160–164.
  35. Liston C, Watts R, Tottenham N, Davidson MC, Niogi S, Ulug AM, Casey BJ (2006): Frontostriatal microstructure modulates efficient recruitment of cognitive control. Cereb Cortex 16:553–560.
  36. Lopez-Lopez C, LeRoith D, Torres-Aleman I (2004): Insulin-like growth factor I is required for vessel remodeling in the adult brain. Proc Natl Acad Sci USA 101:9833–9838.
  37. Ludwig DS (2007): Childhood obesity: the shape of things to come. N Engl J Med 357:2325–2327.
  38. Marais L, Stein DJ, Daniels WM (2009): Exercise increases BDNF levels in the striatum and decreases depressive-like behavior in chronically stressed rats. Metab Brain Dis 24:587–597.
  39. Marques E, Vasconcelos F, Rolo MR, Pereira FC, Silva AP, Macedo TR, Ribeiro CF (2008): Influence of chronic exercise on the amphetamine-induced dopamine release and neurodegeneration in the striatum of the rat. Ann NY Acad Sci 1139:222–231.
  40. Neeper S, Gomez-Pinilla F, Choi J, Cotman CW (1995): Exercise and brain neurotrophins. Nature 373:109.
  41. Oldfield RC (1971): The assessment and analysis of handedness: the Edinburgh inventory. Neuropsychologia 9:97–113.
  42. Olshansky SJ, Passaro DJ, Hershow RC, Layden J, Carnes BA, Brody J, Hayflick L, Butler RN, Allison DB, Ludwig DS (2005): A potential decline in life expectancy in the United States in the 21st century. N Engl J Med 352:1138–1143.
  43. Patenaude B (2007): Bayesian statistical models of shape and appearance for subcortical brain segmentation (D. Phil. thesis). Oxford, University of Oxford.
  44. Patenaude B, Smith SM, Kennedy D, Jenkinson M (2007a): FIRST-FMRIB’s integrated registration and segmentation tool. Chicago, Human Brain Mapping Conference.
  45. Patenaude B, Smith SM, Kennedy D, Jenkinson M (2007b): Bayesian shape and appearance models. Technical report TR07BP1. Oxford, FMRIB Center, University of Oxford.
  46. Ragozzino ME, Jih J, Tzavos A (2002): Involvement of the dorsomedial striatum in behavioral flexibility: role of muscarinic cholinergic receptors. Brain Res 953:205–214.
  47. Shi LH, Luo F, Woodward DJ, Chang JY (2004): Neural responses in multiple basal ganglia regions during spontaneous and treadmill locomotion tasks in rats. Exp Brain Res 157:303–314.
  48. Shvartz E, Reibold RC (1990): Aerobic fitness norms for males and females aged 6 to 75 years: a review. Aviat Space Environ Med 61:3–11.
  49. Sibley BA, Etnier JL (2003): The relationship between physical activity and cognition in children: a meta-analysis. Pediatr Exerc Sci 15:243–256.
  50. Tanner JM (1962): Growth at Adolescence. Oxford, Blackwell Scientific Publications, p 340.
  51. Taylor SJC, Whincup PH, Hindmarsh PC, Lampe F, Odoki K, Cook DG (2001): Performance of a new pubertal self-assessment questionnaire: a preliminary study. Paediatr Perinat Epidemiol 15:88–94.
  52. Tillerson JL, Cohen AD, Philhower J, Miller GW, Zigmond MJ, Schallert T (2001): Forced limb-use effects on the behavioral and neurochemical effects of 6-hydroxydopamine. J Neurosci 21:4427–4435.
  53. Utter AC, Roberson RJ, Nieman DC, Kang J (2002): Children’s OMNI scale of perceived exertion: walking/running evaluation. Med Sci Sports Exerc 34:139–144.
  54. Van Praag H, Christie BR, Sejnowski TJ, Gage FH (1999): Running enhances neurogenesis, learning, and long-term potentiation in mice. Neurobiology 96:13427–13431.
  55. Van Praag H, Shubert T, Zhao C, Gage FH (2005): Exercise enhances learning and hippocampal neurogenesis in aged mice. J Neurosci 25:8680–8685.
  56. Vaynman S, Ying Z, Gomez-Pinilla F (2004): Hippocampal BDNF mediates the efficacy of exercise on synaptic plasticity and cognition. Eur J Neurosci 20:2580–2590.
  57. Wylie SA, van den Wildenberg WP, Ridderinkhof KR, Bashore TR, Powell VD, Manning CA, Wooten GF (2009): The effect of Parkinson’s disease on interference control during action selection. Neuropsychologia 47:145–157.

  

Author Contacts

Laura Chaddock
Department of Psychology, Beckman Institute for Advanced Science and Technology
University of Illinois at Urbana-Champaign
405 North Mathews Avenue, Urbana, IL 61801 (USA)
Tel. +1 610 209 6836, Fax +1 217 333 2922, E-Mail lchaddo2@illinois.edu

  

Article Information

Received: March 29, 2010
Accepted after revision: June 8, 2010
Published online: August 6, 2010
Number of Print Pages : 8
Number of Figures : 1, Number of Tables : 3, Number of References : 57

  

Publication Details

Developmental Neuroscience

Vol. 32, No. 3, Year 2010 (Cover Date: August 2010)

Journal Editor: Levison S.W. (Newark, N.J.)
ISSN: 0378-5866 (Print), eISSN: 1421-9859 (Online)

For additional information: http://www.karger.com/DNE


Copyright / Drug Dosage / Disclaimer

Copyright: All rights reserved. No part of this publication may be translated into other languages, reproduced or utilized in any form or by any means, electronic or mechanical, including photocopying, recording, microcopying, or by any information storage and retrieval system, without permission in writing from the publisher or, in the case of photocopying, direct payment of a specified fee to the Copyright Clearance Center.
Drug Dosage: The authors and the publisher have exerted every effort to ensure that drug selection and dosage set forth in this text are in accord with current recommendations and practice at the time of publication. However, in view of ongoing research, changes in goverment regulations, and the constant flow of information relating to drug therapy and drug reactions, the reader is urged to check the package insert for each drug for any changes in indications and dosage and for added warnings and precautions. This is particularly important when the recommended agent is a new and/or infrequently employed drug.
Disclaimer: The statements, opinions and data contained in this publication are solely those of the individual authors and contributors and not of the publishers and the editor(s). The appearance of advertisements or/and product references in the publication is not a warranty, endorsement, or approval of the products or services advertised or of their effectiveness, quality or safety. The publisher and the editor(s) disclaim responsibility for any injury to persons or property resulting from any ideas, methods, instructions or products referred to in the content or advertisements.

Abstract

The present investigation is the first to explore the association between childhood aerobic fitness and basal ganglia structure and function. Rodent research has revealed that exercise influences the striatum by increasing dopamine signaling and angiogenesis. In children, higher aerobic fitness levels are associated with greater hippocampal volumes, superior performance on tasks of attentional and interference control, and elevated event-related brain potential indices of executive function. The present study used magnetic resonance imaging to investigate if higher-fit and lower-fit 9- and 10-year-old children exhibited differential volumes of other subcortical brain regions, specifically the basal ganglia involved in attentional control. The relationship between aerobic fitness, dorsal and ventral striatum volumes and performance on an attention and inhibition Eriksen flanker task was also examined. The results indicated that higher-fit children showed superior flanker task performance compared to lower-fit children. Higher-fit children also showed greater volumes of the dorsal striatum, and dorsal striatum volume was negatively associated with behavioral interference. The results support the claim that the dorsal striatum is involved in cognitive control and response resolution and that these cognitive processes vary as a function of aerobic fitness. No relationship was found between aerobic fitness, the volume of the ventral striatum and flanker performance. The findings suggest that increased childhood aerobic fitness is associated with greater dorsal striatal volumes and that this is related to enhanced cognitive control. Because children are becoming increasingly overweight, unhealthy and unfit, understanding the neurocognitive benefits of an active lifestyle during childhood has important public health and educational implications.

© 2010 S. Karger AG, Basel


  

Author Contacts

Laura Chaddock
Department of Psychology, Beckman Institute for Advanced Science and Technology
University of Illinois at Urbana-Champaign
405 North Mathews Avenue, Urbana, IL 61801 (USA)
Tel. +1 610 209 6836, Fax +1 217 333 2922, E-Mail lchaddo2@illinois.edu

  

Article Information

Received: March 29, 2010
Accepted after revision: June 8, 2010
Published online: August 6, 2010
Number of Print Pages : 8
Number of Figures : 1, Number of Tables : 3, Number of References : 57

  

Publication Details

Developmental Neuroscience

Vol. 32, No. 3, Year 2010 (Cover Date: August 2010)

Journal Editor: Levison S.W. (Newark, N.J.)
ISSN: 0378-5866 (Print), eISSN: 1421-9859 (Online)

For additional information: http://www.karger.com/DNE


Article / Publication Details

First-Page Preview
Abstract of Original Paper

Received: 3/29/2010
Accepted: 8/6/2010
Published online: 8/6/2010
Issue release date: August 2010

Number of Print Pages: 8
Number of Figures: 1
Number of Tables: 3

ISSN: 0378-5866 (Print)
eISSN: 1421-9859 (Online)

For additional information: http://www.karger.com/DNE


Copyright / Drug Dosage

Copyright: All rights reserved. No part of this publication may be translated into other languages, reproduced or utilized in any form or by any means, electronic or mechanical, including photocopying, recording, microcopying, or by any information storage and retrieval system, without permission in writing from the publisher or, in the case of photocopying, direct payment of a specified fee to the Copyright Clearance Center.
Drug Dosage: The authors and the publisher have exerted every effort to ensure that drug selection and dosage set forth in this text are in accord with current recommendations and practice at the time of publication. However, in view of ongoing research, changes in goverment regulations, and the constant flow of information relating to drug therapy and drug reactions, the reader is urged to check the package insert for each drug for any changes in indications and dosage and for added warnings and precautions. This is particularly important when the recommended agent is a new and/or infrequently employed drug.
Disclaimer: The statements, opinions and data contained in this publication are solely those of the individual authors and contributors and not of the publishers and the editor(s). The appearance of advertisements or/and product references in the publication is not a warranty, endorsement, or approval of the products or services advertised or of their effectiveness, quality or safety. The publisher and the editor(s) disclaim responsibility for any injury to persons or property resulting from any ideas, methods, instructions or products referred to in the content or advertisements.

References

  1. Aguiar AS, Speck AE, Prediger RD, Kapczinski F, Pinho RA (2008): Downhill training upregulates mice hippocampal and striatal brain-derived neurotrophic factor levels. J Neural Transm 115:1251–1255.
  2. American College of Sports Medicine (2006): ACSM’s Guidelines for Exercise Testing and Prescription, ed 7. New York, Lippincott Williams & Wilkins, p 366.
  3. Aron AR, Poldrack RA, Wise SP (2009): Cognition: basal ganglia role; in Squire LR (ed.): Encyclopedia of Neuroscience, vol 2, pp 1069–1077.

    External Resources

  4. Baker JL, Olsen LW, Sorensen TIA (2007): Childhood body-mass index and risk of coronary heart disease in adulthood. N Engl J Med 357:2329–2337.
  5. Bar-Or O (1983): Pediatric sports medicine for the practitioner: From physiologic principles to clinical applications. New York, Springer, p 376.
  6. Birnbaum AS, Lytle LA, Murray DM, Story M, Perry CL, Boutelle KN (2002): Survey development for assessing correlates of young adolescents’ eating. Am J Health Behav 26:284–295.
  7. Buck SM, Hillman CH, Castelli DM (2008): The relation of aerobic fitness to Stroop task performance in preadolescent children. Med Sci Sports Exerc 40:166–172.
  8. Casey BJ, Getz S, Galvan A (2008): The adolescent brain. Dev Rev 28:62–77.
  9. Casey BJ, Thomas KM, Welsh TF, Badgaiyan RD, Eccard CH, Jennings JR, Crone EA (2000): Dissociation of response conflict, attentional selection, and expectancy with functional magnetic resonance imaging. Proc Natl Acad Sci 97:8728–8733.
  10. Casey BJ, Trainor RJ, Orendi JL, Schubert AB, Nystrom LE, Giedd JN, Castellanos FX, Haxby JV, Noll DC, Cohen JD, Forman SD, Dahl RE, Rapoport JL (1997): A developmental functional MRI study of prefrontal activation during performance of a go-no-go task. J Cog Neurosci 9:835–847.
  11. Castelli DM, Hillman CH, Buck SM, Erwin HE (2007): Physical fitness and academic achievement in third- and fifth-grade students. J Sport Exerc Psychol 29:239–252.
  12. Chaddock L, Erickson KI, Prakash RS, Kim JS, Voss MW, VanPatter M, Pontifex MB, Raine LB, Konkel A, Hillman CH, Cohen NJ, Kramer AF (2010): A neuroimaging investigation of the association between aerobic fitness, hippocampal volume and memory performance in preadolescent children. Devel- opment & Aging. Abstr Cogn Neurosci Soc B82-76.
  13. Chaddock L, Hillman CH, Buck SM, Cohen NJ (in press): Aerobic fitness and executive control of relational memory in preadolescent children: MSSE.
  14. Chomitz VR, Slining MM, McGowan RJ, Mitchell SE, Dawson GF, Hacker KA (2009): Is there a relationship between physical fitness and academic achievement? Positive results from public school children in the northeastern United States. J Sch Health 79:30–37.
  15. Colcombe S, Kramer AF (2003): Fitness effects on the cognitive function of older adults: A meta-analytic study. Psychol Sci 14:125–130.
  16. Colcombe SJ, Kramer AF, Erickson KI, Scalf P, McAuley E, Cohen NJ, Webb A, Jerome GJ, Marquez DX, Elavsky S (2004): Cardiovascular fitness, cortical plasticity, and aging. PNAS 101:3316–3321.
  17. Cotman CW, Berchtold NC (2002): Exercise: a behavioral intervention to enhance brain health and plasticity. Trends Neurosci 25:295–301.
  18. Di Martino A, Scheres A, Marguiles DS, Kelly AM, Uddin LQ, Shehzad Z, Biswal B, Walters JR, Castellanos FX, Milham MP (2008): Functional connectivity of human striatum: a resting state fMRI study. Cereb Cortex 18:2735–2747.
  19. Ding YH, Luan XD, Li J, Rafols JA, Guthinkonda M, Diaz FG, Ding Y (2004): Exercise-induced overexpression of angiogenic factors and reduction of ischemia/reperfusion injury in stroke. Curr Neurovasc Res 1:411–420.
  20. Draganski B, Kherif F, Kloppel S, Cook PA, Alexander DC, Parker GJ, Deichmann R, Ashburner J, Frackowiak RS (2008): Evidence for segregated and integrative connectivity patterns in the human basal ganglia. J Neurosci 28:7143–7152.
  21. DuPaul GJ, Power TJ, Anastopoulos A, Reid R (1998): ADHD Rating Scale – IV: Checklists, Norms, and Clinical Interpretation. New York, Guilford Press.
  22. Erickson KI, Boot WR, Basak C, Neider MR, Prakash RS, Voss MW, Graybiel AM, Simons DJ, Fabiani M, Gratton G, Kramer AF (in press): Striatal volume predicts level of video game skill acquisition.
  23. Erickson KI, Prakash RS, Voss MW, Chaddock L, Hu L, Morris KS, White SM, Wojcicki TR, McAuley E, Kramer AF (2009): Aerobic fitness is associated with hippocampal volume in elderly humans. Hippocampus 19:1030–1039.
  24. Eriksen BA, Eriksen CW (1974): Effects of noise letters on the identification of a target letter in a nonsearch task. Percept Psychophys 16:143–149.
  25. Freedson PS, Goodman TL (1993): Measurement of oxygen consumption; in Rowland TW (ed): Pediatric Laboratory Exercise Testing: Clinical Guidelines. Human Kinetics, Champaign, pp 91–113.
  26. Graybiel AM (2005): The basal ganglia: learning new tricks and loving it. Curr Opin Neurobiol 15:638–644.
  27. Graybiel AM (2008): Habits, rituals and the evaluative brain. Annu Rev Neurosci 31:359–387.
  28. Hillman CH, Buck SM, Themanson JR, Pontifex MB, Castelli DM (2009): Aerobic fitness and cognitive development: event-related brain potential and task performance of executive control in preadolescent children. Dev Psychol 45:114–129.
  29. Hillman CH, Castelli DM, Buck SM (2005): Aerobic fitness and neurocognitive function in healthy preadolescent children. Med Sci Sports Exerc 37:1967–1974.
  30. Hillman CH, Erickson KI, Kramer AF (2008): Be smart, exercise your heart: exercise effects on brain and cognition. Nature Rev Neurosci 9:58–65.
  31. Kaufman AS, Kaufman NL (1990): Kaufman Brief Intelligence Test. Circle Pines, AGS.
  32. Kramer AF, Hahn S, Cohen N, Banich M, McAuley E, Harrison C, Chason J, Vakil E, Bardell L, Boileau RA, Colcombe A (1999): Aging, fitness, and neurocognitive function. Nature 400:418–419.
  33. Kramer A, Humphrey D, Larish J, Logan G, Strayer D (1994): Aging and inhibition: beyond a unitary view of inhibitory processing in attention. Psychol Aging 9:491–512.
  34. Li J, Ding YH, Rafols JA, Lai Q, McAllister JP, Ding Y (2005): Increased astrocyte proliferation in rats after running exercise. Neurosci Lett 386:160–164.
  35. Liston C, Watts R, Tottenham N, Davidson MC, Niogi S, Ulug AM, Casey BJ (2006): Frontostriatal microstructure modulates efficient recruitment of cognitive control. Cereb Cortex 16:553–560.
  36. Lopez-Lopez C, LeRoith D, Torres-Aleman I (2004): Insulin-like growth factor I is required for vessel remodeling in the adult brain. Proc Natl Acad Sci USA 101:9833–9838.
  37. Ludwig DS (2007): Childhood obesity: the shape of things to come. N Engl J Med 357:2325–2327.
  38. Marais L, Stein DJ, Daniels WM (2009): Exercise increases BDNF levels in the striatum and decreases depressive-like behavior in chronically stressed rats. Metab Brain Dis 24:587–597.
  39. Marques E, Vasconcelos F, Rolo MR, Pereira FC, Silva AP, Macedo TR, Ribeiro CF (2008): Influence of chronic exercise on the amphetamine-induced dopamine release and neurodegeneration in the striatum of the rat. Ann NY Acad Sci 1139:222–231.
  40. Neeper S, Gomez-Pinilla F, Choi J, Cotman CW (1995): Exercise and brain neurotrophins. Nature 373:109.
  41. Oldfield RC (1971): The assessment and analysis of handedness: the Edinburgh inventory. Neuropsychologia 9:97–113.
  42. Olshansky SJ, Passaro DJ, Hershow RC, Layden J, Carnes BA, Brody J, Hayflick L, Butler RN, Allison DB, Ludwig DS (2005): A potential decline in life expectancy in the United States in the 21st century. N Engl J Med 352:1138–1143.
  43. Patenaude B (2007): Bayesian statistical models of shape and appearance for subcortical brain segmentation (D. Phil. thesis). Oxford, University of Oxford.
  44. Patenaude B, Smith SM, Kennedy D, Jenkinson M (2007a): FIRST-FMRIB’s integrated registration and segmentation tool. Chicago, Human Brain Mapping Conference.
  45. Patenaude B, Smith SM, Kennedy D, Jenkinson M (2007b): Bayesian shape and appearance models. Technical report TR07BP1. Oxford, FMRIB Center, University of Oxford.
  46. Ragozzino ME, Jih J, Tzavos A (2002): Involvement of the dorsomedial striatum in behavioral flexibility: role of muscarinic cholinergic receptors. Brain Res 953:205–214.
  47. Shi LH, Luo F, Woodward DJ, Chang JY (2004): Neural responses in multiple basal ganglia regions during spontaneous and treadmill locomotion tasks in rats. Exp Brain Res 157:303–314.
  48. Shvartz E, Reibold RC (1990): Aerobic fitness norms for males and females aged 6 to 75 years: a review. Aviat Space Environ Med 61:3–11.
  49. Sibley BA, Etnier JL (2003): The relationship between physical activity and cognition in children: a meta-analysis. Pediatr Exerc Sci 15:243–256.
  50. Tanner JM (1962): Growth at Adolescence. Oxford, Blackwell Scientific Publications, p 340.
  51. Taylor SJC, Whincup PH, Hindmarsh PC, Lampe F, Odoki K, Cook DG (2001): Performance of a new pubertal self-assessment questionnaire: a preliminary study. Paediatr Perinat Epidemiol 15:88–94.
  52. Tillerson JL, Cohen AD, Philhower J, Miller GW, Zigmond MJ, Schallert T (2001): Forced limb-use effects on the behavioral and neurochemical effects of 6-hydroxydopamine. J Neurosci 21:4427–4435.
  53. Utter AC, Roberson RJ, Nieman DC, Kang J (2002): Children’s OMNI scale of perceived exertion: walking/running evaluation. Med Sci Sports Exerc 34:139–144.
  54. Van Praag H, Christie BR, Sejnowski TJ, Gage FH (1999): Running enhances neurogenesis, learning, and long-term potentiation in mice. Neurobiology 96:13427–13431.
  55. Van Praag H, Shubert T, Zhao C, Gage FH (2005): Exercise enhances learning and hippocampal neurogenesis in aged mice. J Neurosci 25:8680–8685.
  56. Vaynman S, Ying Z, Gomez-Pinilla F (2004): Hippocampal BDNF mediates the efficacy of exercise on synaptic plasticity and cognition. Eur J Neurosci 20:2580–2590.
  57. Wylie SA, van den Wildenberg WP, Ridderinkhof KR, Bashore TR, Powell VD, Manning CA, Wooten GF (2009): The effect of Parkinson’s disease on interference control during action selection. Neuropsychologia 47:145–157.