Sports Medicine

, Volume 43, Issue 5, pp 301–311 | Cite as

Neurophysiological Determinants of Theoretical Concepts and Mechanisms Involved in Pacing

  • Bart Roelands
  • Jos de Koning
  • Carl Foster
  • Floor Hettinga
  • Romain MeeusenEmail author
Review Article


Fatigue during prolonged exercise is often described as an acute impairment of exercise performance that leads to an inability to produce or maintain a desired power output. In the past few decades, interest in how athletes experience fatigue during competition has grown enormously. Research has evolved from a dominant focus on peripheral causes of fatigue towards a complex interplay between peripheral and central limitations of performance. Apparently, both feedforward and feedback mechanisms, based on the principle of teleoanticipation, regulate power output (e.g. speed) during a performance. This concept is called ‘pacing’ and represents the use of energetic resources during exercise, in a way such that all energy stores are used before finishing a race, but not so far from the end of a race that a meaningful slowdown can occur.

It is believed that the pacing selected by athletes is largely dependent on the anticipated exercise duration and on the presence of an experientially developed performance template. Most studies investigating pacing during prolonged exercise in ambient temperatures, have observed a fast start, followed by an even pace strategy in the middle of the event with an end sprint in the final minutes of the race. A reduction in pace observed at commencement of the event is often more evident during exercise in hot environmental conditions. Further, reductions in power output and muscle activation occur before critical core temperatures are reached, indicating that subjects can anticipate the exercise intensity and heat stress they will be exposed to, resulting in a tactical adjustment of the power output. Recent research has shown that not only climatic stress but also pharmacological manipulation of the central nervous system has the ability to cause changes in endurance performance. Subjects seem to adapt their strategy specifically in the early phases of an exercise task. In high-ambient temperatures, dopaminergic manipulations clearly improve performance. The distribution of the power output reveals that after dopamine reuptake inhibition, subjects are able to maintain a higher power output compared with placebo. Manipulations of serotonin and, especially, noradrenaline, have the opposite effect and force subjects to decrease power output early in the time trial. Interestingly, after manipulation of brain serotonin, subjects are often unable to perform an end sprint, indicating an absence of a reserve capacity or motivation to increase power output. Taken together, it appears that many factors, such as ambient conditions and manipulation of brain neurotransmitters, have the potential to influence power output during exercise, and might thus be involved as regulatory mechanisms in the complex skill of pacing.


Power Output Time Trial Reboxetine Pace Strategy Central Nervous System Drug 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.



Funding: Bart Roelands is a postdoctoral fellow of the Research Fund of Flanders (FWO). We want to acknowledge funding through the Vrije Universiteit Brussel (OZR 607, 990, 1235).

Conflict of interest

The authors have no conflicts of interest that are directly relevant to the content of this review.


  1. 1.
    Meeusen R, Watson P, Hasegawa H, Roelands B, Piacentini MF. Central fatigue: the serotonin hypothesis and beyond. Sports Med. 2006;36(10):881–909.PubMedCrossRefGoogle Scholar
  2. 2.
    Amann M. Central and peripheral fatigue: interaction during cycling exercise in humans. Med Sci Sports Exerc. 2011;43(11):2039–45.PubMedCrossRefGoogle Scholar
  3. 3.
    Roelands B, Meeusen R. Alterations in central fatigue by pharmacological manipulations of neurotransmitters in normal and high ambient temperature. Sports Med. 2010;40(3):229–46.PubMedCrossRefGoogle Scholar
  4. 4.
    St Clair Gibson A, Lambert EV, Rauch LH, Tucker R, Baden DA, Foster C, et al. The role of information processing between the brain and peripheral physiological systems in pacing and perception of effort. Sports Med. 2006;36(8):705–22.PubMedCrossRefGoogle Scholar
  5. 5.
    Tucker R, Rauch L, Harley YX, Noakes TD. Impaired exercise performance in the heat is associated with an anticipatory reduction in skeletal muscle recruitment. Pflugers Arch. 2004;448(4):422–30.PubMedCrossRefGoogle Scholar
  6. 6.
    Robinson S, Robinson DL, Mountjoy RJ, Bullard RW. Influence of fatigue on the efficiency of men during exhausting runs. J Appl Physiol. 1958;12(2):197–201.PubMedGoogle Scholar
  7. 7.
    Foster C, Snyder AC, Thompson NN, Green MA, Foley M, Schrager M. Effect of pacing strategy on cycle time trial performance. Med Sci Sports Exerc. 1993;25(3):383–8.PubMedGoogle Scholar
  8. 8.
    Foster C, Schrager M, Snyder AC, Thompson NN. Pacing strategy and athletic performance. Sports Med. 1994;17(2):77–85.PubMedCrossRefGoogle Scholar
  9. 9.
    Foster C, de Koning J, BIschel S, Casolino E, Malterer K, O’Brien K, et al. Pacing Strategies for Endurance Performance. In: Mujika I, editor. Endurance training, science and practice. 1 ed. Vitoria-Gasteiz: Inigo Mujika S.L.U.; 2012.Google Scholar
  10. 10.
    Tucker R, Noakes TD. The physiological regulation of pacing strategy during exercise: a critical review. Br J Sports Med. 2009;43(6):e1.PubMedCrossRefGoogle Scholar
  11. 11.
    Abbiss CR, Laursen PB. Describing and understanding pacing strategies during athletic competition. Sports Med. 2008;38(3):239–52.PubMedCrossRefGoogle Scholar
  12. 12.
    de Koning JJ, Foster C, Bakkum A, Kloppenburg S, Thiel C, Joseph T, et al. Regulation of pacing strategy during athletic competition. PLoS One. 2011;6(1):e15863.PubMedCrossRefGoogle Scholar
  13. 13.
    Paterson S, Marino FE. Effect of deception of distance on prolonged cycling performance. Percept Mot Skills. 2004;98(3 Pt 1):1017–26.PubMedCrossRefGoogle Scholar
  14. 14.
    St Clair Gibson A, Goedecke JH, Harley YX, Myers LJ, Lambert MI, Noakes TD, et al. Metabolic setpoint control mechanisms in different physiological systems at rest and during exercise. J Theor Biol. 2005;236(1):60–72.PubMedCrossRefGoogle Scholar
  15. 15.
    Borg GA. Psychophysical bases of perceived exertion. Med Sci Sports Exerc. 1982;14(5):377–81.PubMedGoogle Scholar
  16. 16.
    Hampson DB, St Clair Gibson A, Lambert MI, Noakes TD. The influence of sensory cues on the perception of exertion during exercise and central regulation of exercise performance. Sports Med. 2001;31(13):935–52.PubMedCrossRefGoogle Scholar
  17. 17.
    St Clair Gibson A, Noakes TD. Evidence for complex system integration and dynamic neural regulation of skeletal muscle recruitment during exercise in humans. Br J Sports Med. 2004;38(6):797–806.PubMedCrossRefGoogle Scholar
  18. 18.
    Craig AD. Interoception: the sense of the physiological condition of the body. Curr Opin Neurobiol. 2003;13(4):500–5.PubMedCrossRefGoogle Scholar
  19. 19.
    Hettinga FJ, de Koning JJ, Hulleman M, Foster C. Relative importance of pacing strategy and mean power output in 1500-m self-paced cycling. Br J Sports Med. 2012;46(1):30–5.PubMedCrossRefGoogle Scholar
  20. 20.
    Tucker R. The anticipatory regulation of performance: the physiological basis for pacing strategies and the development of a perception-based model for exercise performance. Br J Sports Med. 2009;43(6):392–400.PubMedCrossRefGoogle Scholar
  21. 21.
    Abbiss CR, Laursen PB. Models to explain fatigue during prolonged endurance cycling. Sports Med. 2005;35(10):865–98.PubMedCrossRefGoogle Scholar
  22. 22.
    Hill AV. Muscular activity and carbohydrate metabolism. Science. 1924;60(1562):505–14.PubMedCrossRefGoogle Scholar
  23. 23.
    Ulmer HV. Concept of an extracellular regulation of muscular metabolic rate during heavy exercise in humans by psychophysiological feedback. Experientia. 1996;52(5):416–20.PubMedCrossRefGoogle Scholar
  24. 24.
    Noakes TD, St Clair Gibson A, Lambert EV. From catastrophe to complexity: a novel model of integrative central neural regulation of effort and fatigue during exercise in humans: summary and conclusions. Br J Sports Med. 2005;39(2):120–4.PubMedCrossRefGoogle Scholar
  25. 25.
    Noakes TD. 1996 J.B. Wolffe Memorial Lecture. Challenging beliefs: ex Africa semper aliquid novi. Med Sci Sports Exerc. 1997;29(5):571–90.PubMedCrossRefGoogle Scholar
  26. 26.
    Reid C. The mechanism of voluntary muscular fatigue. Br Med J. 1927;2(3481):545–6.PubMedCrossRefGoogle Scholar
  27. 27.
    Swart J, Lamberts RP, Lambert MI, Gibson ASC, Lambert EV, Skowno J, et al. Exercising with reserve: evidence that the central nervous system regulates prolonged exercise performance. Br J Sports Med. 2009;43(10):782–8.PubMedCrossRefGoogle Scholar
  28. 28.
    Kaufman MP, Hayes SG, Adreani CM, Pickar JG. Discharge properties of group III and IV muscle afferents. Adv Exp Med Biol. 2002;508:25–32.PubMedCrossRefGoogle Scholar
  29. 29.
    Amann M, Proctor LT, Sebranek JJ, Eldridge MW, Pegelow DF, Dempsey JA. Somatosensory feedback from the limbs exerts inhibitory influences on central neural drive during whole body endurance exercise. J Appl Physiol. 2008;105(6):1714–24.PubMedCrossRefGoogle Scholar
  30. 30.
    Amann M, Dempsey JA. Locomotor muscle fatigue modifies central motor drive in healthy humans and imposes a limitation to exercise performance. J Physiol. 2008;586(1):161–73.PubMedCrossRefGoogle Scholar
  31. 31.
    Amann M, Proctor LT, Sebranek JJ, Pegelow DF, Dempsey JA. Opioid-mediated muscle afferents inhibit central motor drive and limit peripheral muscle fatigue development in humans. J Physiol. 2009;587(Pt 1):271–83.PubMedCrossRefGoogle Scholar
  32. 32.
    Lambert EV, St Clair Gibson A, Noakes TD. Complex systems model of fatigue: integrative homoeostatic control of peripheral physiological systems during exercise in humans. Br J Sports Med. 2005;39(1):52–62.PubMedCrossRefGoogle Scholar
  33. 33.
    de Koning JJ, Bobbert MF, Foster C. Determination of optimal pacing strategy in track cycling with an energy flow model. J Sci Med Sport. 1999;2(3):266–77.PubMedCrossRefGoogle Scholar
  34. 34.
    Thomas K, Stone MR, Thompson KG, St Clair Gibson A, Ansley L. Reproducibility of pacing strategy during simulated 20-km cycling time trials in well-trained cyclists. Eur J Appl Physiol. 2012;112(1):223–9.PubMedCrossRefGoogle Scholar
  35. 35.
    Renfree A, West J, Corbett M, Rhoden C, St Clair Gibson A. Complex interplay between determinants of pacing and performance during 20 km cycle time trials. Int J Sports Physiol Perform. 2012;7:121–9.PubMedGoogle Scholar
  36. 36.
    Garland SW. An analysis of the pacing strategy adopted by elite competitors in 2000 m rowing. Br J Sports Med. 2005;39(1):39–42.PubMedCrossRefGoogle Scholar
  37. 37.
    Lima-Silva AE, Bertuzzi RC, Pires FO, Barros RV, Gagliardi JF, Hammond J, et al. Effect of performance level on pacing strategy during a 10-km running race. Eur J Appl Physiol. 2010;108(5):1045–53.PubMedCrossRefGoogle Scholar
  38. 38.
    Mattern CO, Kenefick RW, Kertzer R, Quinn TJ. Impact of starting strategy on cycling performance. Int J Sports Med. 2001;22(5):350–5.PubMedCrossRefGoogle Scholar
  39. 39.
    Hausswirth C, Le Meur Y, Bieuzen F, Brisswalter J, Bernard T. Pacing strategy during the initial phase of the run in triathlon: influence on overall performance. Eur J Appl Physiol. 2010;108(6):1115–23.PubMedCrossRefGoogle Scholar
  40. 40.
    Hettinga FJ, De Koning JJ, Schmidt LJ, Wind NA, Macintosh BR, Foster C. Optimal pacing strategy: from theoretical modelling to reality in 1500-m speed skating. Br J Sports Med. 2011;45(1):30–5.PubMedCrossRefGoogle Scholar
  41. 41.
    Watson P, Hasegawa H, Roelands B, Piacentini MF, Looverie R, Meeusen R. Acute dopamine/noradrenaline reuptake inhibition enhances human exercise performance in warm, but not temperate conditions. J Physiol. 2005;565(Pt 3):873–83.PubMedCrossRefGoogle Scholar
  42. 42.
    Roelands B, Hasegawa H, Watson P, Piacentini MF, Buyse L, De Schutter G, et al. The effects of acute dopamine reuptake inhibition on performance. Med Sci Sports Exerc. 2008;40(5):879–85.PubMedCrossRefGoogle Scholar
  43. 43.
    Roelands B, Goekint M, Heyman E, Piacentini MF, Watson P, Hasegawa H, et al. Acute norepinephrine reuptake inhibition decreases performance in normal and high ambient temperature. J Appl Physiol. 2008;105(1):206–12.PubMedCrossRefGoogle Scholar
  44. 44.
    Roelands B, Goekint M, Buyse L, Pauwels F, De Schutter G, Piacentini F, et al. Time trial performance in normal and high ambient temperature: is there a role for 5-HT? Eur J Appl Physiol. 2009;107(1):119–26.PubMedCrossRefGoogle Scholar
  45. 45.
    Roelands B, Hasegawa H, Watson P, Piacentini MF, Buyse L, De Schutter G, et al. Performance and thermoregulatory effects of chronic bupropion administration in the heat. Eur J Appl Physiol. 2009;105(3):493–8.PubMedCrossRefGoogle Scholar
  46. 46.
    Roelands B, Buyse L, Pauwels F, Delbeke F, Deventer K, Meeusen R. No effect of caffeine on exercise performance in high ambient temperature. Eur J Appl Physiol. 2011;111(12):3089–95.PubMedCrossRefGoogle Scholar
  47. 47.
    Galloway SD, Maughan RJ. Effects of ambient temperature on the capacity to perform prolonged cycle exercise in man. Med Sci Sports Exerc. 1997;29(9):1240–9.PubMedCrossRefGoogle Scholar
  48. 48.
    Parkin JM, Carey MF, Zhao S, Febbraio MA. Effect of ambient temperature on human skeletal muscle metabolism during fatiguing submaximal exercise. J Appl Physiol. 1999;86(3):902–8.PubMedGoogle Scholar
  49. 49.
    Tatterson AJ, Hahn AG, Martin DT, Febbraio MA. Effects of heat stress on physiological responses and exercise performance in elite cyclists. J Sci Med Sport. 2000;3(2):186–93.PubMedCrossRefGoogle Scholar
  50. 50.
    Jeukendrup A, Saris WH, Brouns F, Kester AD. A new validated endurance performance test. Med Sci Sports Exerc. 1996;28(2):266–70.PubMedCrossRefGoogle Scholar
  51. 51.
    Hickey MS, Costill DL, McConell GK, Widrick JJ, Tanaka H. Day to day variation in time trial cycling performance. Int J Sports Med. 1992;13(6):467–70.PubMedCrossRefGoogle Scholar
  52. 52.
    Klass M, Roelands B, Levenez M, Fontenelle V, Pattyn N, Meeusen R, et al. Effects of noradrenaline and dopamine on supraspinal fatigue in well-trained men. Med Sci Sports Exerc. 2012;44(12):2299–308.PubMedCrossRefGoogle Scholar
  53. 53.
    Roelands B, Watson P, Cordery P, Decoster S, Debaste E, Maughan R, et al. A dopamine/noradrenaline reuptake inhibitor improves performance in the heat, but only at the maximum therapeutic dose. Scand J Med Sci Sports. 2012;22(5):e93–8.PubMedCrossRefGoogle Scholar
  54. 54.
    Cheung SS. Hyperthermia and voluntary exhaustion: integrating models and future challenges. Appl Physiol Nutr Metab. 2007;32(4):808–17.PubMedCrossRefGoogle Scholar
  55. 55.
    Gonzalez-Alonso J, Teller C, Andersen SL, Jensen FB, Hyldig T, Nielsen B. Influence of body temperature on the development of fatigue during prolonged exercise in the heat. J Appl Physiol. 1999;86(3):1032–9.PubMedGoogle Scholar
  56. 56.
    Walters TJ, Ryan KL, Tate LM, Mason PA. Exercise in the heat is limited by a critical internal temperature. J Appl Physiol. 2000;89(2):799–806.PubMedGoogle Scholar
  57. 57.
    Nybo L, Nielsen B. Hyperthermia and central fatigue during prolonged exercise in humans. J Appl Physiol. 2001;91(3):1055–60.PubMedGoogle Scholar
  58. 58.
    Fuller A, Carter RN, Mitchell D. Brain and abdominal temperatures at fatigue in rats exercising in the heat. J Appl Physiol. 1998;84(3):877–83.PubMedGoogle Scholar
  59. 59.
    Abbiss CR, Burnett A, Nosaka K, Green JP, Foster JK, Laursen PB. Effect of hot versus cold climates on power output, muscle activation, and perceived fatigue during a dynamic 100-km cycling trial. J Sports Sci. 2010;28(2):117–25.PubMedCrossRefGoogle Scholar
  60. 60.
    Ely BR, Cheuvront SN, Kenefick RW, Sawka MN. Aerobic performance is degraded, despite modest hyperthermia, hot environments. Med Sci Sports Exerc. 2010;42(1):135–41.PubMedCrossRefGoogle Scholar
  61. 61.
    Abbiss C, Peiffer J, Wall B, Martin D, Laursen P. Influence of starting strategy on cycling time trial performance in the heat. Int J Sports Med. 2009;30(03):188–93.PubMedCrossRefGoogle Scholar
  62. 62.
    Barwood MJ, Corbett J, White D, James J. Early change in thermal perception is not a driver of anticipatory exercise pacing in the heat. Br J Sports Med. 2012;46:936–42.PubMedCrossRefGoogle Scholar
  63. 63.
    Levels K, de Koning JJ, Foster C, Daanen HA. The effect of skin temperature on performance during a 7.5-km cycling time trial. Eur J Appl Physiol. 2012;112:3387–95.PubMedCrossRefGoogle Scholar
  64. 64.
    Crewe H, Tucker R, Noakes TD. The rate of increase in rating of perceived exertion predicts the duration of exercise to fatigue at a fixed power output in different environmental conditions. Eur J Appl Physiol. 2008;103(5):569–77.PubMedCrossRefGoogle Scholar
  65. 65.
    Tucker R, Marle T, Lambert EV, Noakes TD. The rate of heat storage mediates an anticipatory reduction in exercise intensity during cycling at a fixed rating of perceived exertion. J Physiol. 2006;574(Pt 3):905–15.PubMedCrossRefGoogle Scholar
  66. 66.
    Skein M, Duffield R. The effects of fluid ingestion on free-paced intermittent-sprint performance and pacing strategies in the heat. J Sports Sci. 2010;28(3):299–307.PubMedCrossRefGoogle Scholar
  67. 67.
    Corbett J, Barwood MJ, Ouzounoglou A, Thelwell R, Dicks M. Influence of competition on performance and pacing during cycling exercise. Med Sci Sports Exerc. 2012;44(3):509–15.PubMedCrossRefGoogle Scholar
  68. 68.
    Impellizzeri FM. Psychobiological factors are more important than central fatigue in limiting endurance performance. J Appl Physiol. 2010;108(2):459 (author reply 69).Google Scholar
  69. 69.
    Meeusen R, Roelands B. Central fatigue and neurotransmitters, can thermoregulation be manipulated? Scand J Med Sci Sports. 2010;20(Suppl 3):19–28.PubMedCrossRefGoogle Scholar
  70. 70.
    Bridge MW, Weller AS, Rayson M, Jones DA. Responses to exercise in the heat related to measures of hypothalamic serotonergic and dopaminergic function. Eur J Appl Physiol. 2003;89(5):451–9.PubMedCrossRefGoogle Scholar
  71. 71.
    Cooper BR, Wang CM, Cox RF, Norton R, Shea V, Ferris RM. Evidence that the acute behavioral and electrophysiological effects of bupropion (Wellbutrin) are mediated by a noradrenergic mechanism. Neuropsychopharmacology. 1994;11(2):133–41.PubMedGoogle Scholar
  72. 72.
    Piacentini MF, Meeusen R, Buyse L, De Schutter G, Kempenaers F, Van Nijvel J, et al. No effect of a noradrenergic reuptake inhibitor on performance in trained cyclists. Med Sci Sports Exerc. 2002;34(7):1189–93.PubMedCrossRefGoogle Scholar

Copyright information

© Springer International Publishing Switzerland 2013

Authors and Affiliations

  • Bart Roelands
    • 1
    • 2
  • Jos de Koning
    • 3
    • 4
  • Carl Foster
    • 3
    • 4
  • Floor Hettinga
    • 5
  • Romain Meeusen
    • 1
    Email author
  1. 1.Department of Human Physiology, Faculty of Physical Education and PhysiotherapyVrije Universiteit BrusselBrusselsBelgium
  2. 2.Fund for Scientific Research Flanders (FWO)BrusselsBelgium
  3. 3.Research Institute MOVE, Faculty of Human Movement SciencesVU UniversityAmsterdamThe Netherlands
  4. 4.Department of Exercise and Sport ScienceUniversity of WisconsinLa CrosseUSA
  5. 5.University of GroningenUniversity Medical Center Groningen, Center of Human Movement SciencesGroningenThe Netherlands

Personalised recommendations