Sports Medicine

, Volume 39, Issue 3, pp 179–206 | Cite as

Physiological Differences Between Cycling and Running

Lessons from Triathletes
  • Gregoire P. Millet
  • V. E. Vleck
  • D. J. Bentley
Review Article

Abstract

The purpose of this review was to provide a synopsis of the literature concerning the physiological differences between cycling and running. By comparing physiological variables such as maximal oxygen consumption (V̇O2max), anaerobic threshold (AT), heart rate, economy or delta efficiency measured in cycling and running in triathletes, runners or cyclists, this review aims to identify the effects of exercise modality on the underlying mechanisms (ventilatory responses, blood flow,muscle oxidative capacity, peripheral innervation and neuromuscular fatigue) of adaptation. The majority of studies indicate that runners achieve a higher V̇O2max on treadmill whereas cyclists can achieve a V̇O2max value in cycle ergometry similar to that in treadmill running. Hence, V̇O2max is specific to the exercise modality. In addition, the muscles adapt specifically to a given exercise task over a period of time, resulting in an improvement in submaximal physiological variables such as the ventilatory threshold, in some cases without a change in V̇O2max. However, this effect is probably larger in cycling than in running. At the same time, skill influencing motor unit recruitment patterns is an important influence on the anaerobic threshold in cycling. Furthermore, it is likely that there is more physiological training transfer from running to cycling than vice versa. In triathletes, there is generally no difference in V̇O2max measured in cycle ergometry and treadmill running. The data concerning the anaerobic threshold in cycling and running in triathletes are conflicting. This is likely to be due to a combination of actual training load and prior training history in each discipline. The mechanisms surrounding the differences in the AT together with V̇O2max in cycling and running are not largely understood but are probably due to the relative adaptation of cardiac output influencing V̇O2max and also the recruitment of muscle mass in combination with the oxidative capacity of this mass influencing the AT. Several other physiological differences between cycling and running are addressed: heart rate is different between the two activities both for maximal and submaximal intensities. The delta efficiency is higher in running. Ventilation is more impaired in cycling than in running. It has also been shown that pedalling cadence affects the metabolic responses during cycling but also during a subsequent running bout. However, the optimal cadence is still debated. Central fatigue and decrease in maximal strength are more important after prolonged exercise in running than in cycling.

References

  1. 1.
    Joyner MJ, Coyle EF. Endurance exercise performance: the physiology of champions. J Physiol 2008 Jan 1; 586 (1): 35–44PubMedCrossRefGoogle Scholar
  2. 2.
    Loy SF, Hoffmann JJ, Holland GJ. Benefits and practical use of cross-training in sports. Sports Med 1995 Jan; 19(1): 1–8PubMedCrossRefGoogle Scholar
  3. 3.
    Tanaka H. Effects of cross-training: transfer of training effects on VO2max between cycling, running and swimming. Sports Med 1994 Nov; 18 (5): 330–9PubMedCrossRefGoogle Scholar
  4. 4.
    Sleivert GG, Rowlands DS. Physical and physiological factors associated with success in the triathlon. Sports Med 1996 Jul; 22 (1): 8–18PubMedCrossRefGoogle Scholar
  5. 5.
    Pechar GS, McArdle WD, Katch FI, et al. Specificity of cardiorespiratory adaptation to bicycle and treadmill training. J Appl Physiol 1974 Jun; 36 (6): 753–6PubMedGoogle Scholar
  6. 6.
    Withers RT, Sherman WM, Miller JM, et al. Specificity of the anaerobic threshold in endurance trained cyclists and runners. Eur J Appl Physiol Occup Physiol 1981; 47 (1): 93–104PubMedCrossRefGoogle Scholar
  7. 7.
    Fernhall B, Kohrt W. The effect of training specificity on maximal and submaximal physiological responses to treadmill and cycle ergometry. J Sports Med Phys Fitness 1990 Sep; 30 (3): 268–75PubMedGoogle Scholar
  8. 8.
    Basset FA, Boulay MR. Specificity of treadmill and cycle ergometer tests in triathletes, runners and cyclists. Eur J Appl Physiol 2000 Feb; 81 (3): 214–21PubMedCrossRefGoogle Scholar
  9. 9.
    Hue O, Le Gallais D, Chollet D, et al. Ventilatory threshold and maximal oxygen uptake in present triathletes. Can J Appl Physiol 2000 Apr; 25 (2): 102–13PubMedCrossRefGoogle Scholar
  10. 10.
    Schneider DA, Lacroix KA, Atkinson GR, et al. Ventilatory threshold and maximal oxygen uptake during cycling and running in triathletes. Med Sci Sports Exerc 1990 Apr; 22 (2): 257–64PubMedGoogle Scholar
  11. 11.
    Millet GP, Dreano P, Bentley DJ. Physiological characteristics of elite short- and long-distance triathletes. Eur J Appl Physiol 2003 Jan; 88 (4-5): 427–30PubMedCrossRefGoogle Scholar
  12. 12.
    Kreider RB. Ventilatory threshold in swimming, cycling and running in triathletes. Int J Sports Med 1988; 9: 147–8Google Scholar
  13. 13.
    Millet GP, Candau RB, Barbier B, et al. Modelling the transfers of training effects on performance in elite triathletes. Int J Sports Med 2002 Jan; 23 (1): 55–63PubMedCrossRefGoogle Scholar
  14. 14.
    Astrand PO, Saltin B. Maximal oxygen uptake and heart rate in various types of muscular activity. J Appl Physiol 1961 Nov; 16: 977–81PubMedGoogle Scholar
  15. 15.
    Saltin B. The interplay between peripheral and central factors in the adaptive response to exercise and training. Ann N Y Acad Sci 1977; 301: 224–31PubMedCrossRefGoogle Scholar
  16. 16.
    Saltin B, Nazar K, Costill DL, et al. The nature of the training response; peripheral and central adaptations ofone-legged exercise. Acta Physiol Scand 1976 Mar; 96 (3): 289–305PubMedCrossRefGoogle Scholar
  17. 17.
    Gleser MA, Horstman DH, Mello RP. The effect on VO2max of adding arm work to maximal leg work. Med Sci Sports 1974 Summer; 6 (2): 104–7PubMedGoogle Scholar
  18. 18.
    Secher NH, Ruberg-Larsen N, Binkhorst RA, et al. Maximal oxygen uptake during arm cranking and combined arm plus leg exercise. J Appl Physiol 1974 May; 36 (5): 515–8PubMedGoogle Scholar
  19. 19.
    Reybrouck T, Heigenhauser GF, Faulkner JA. Limitations to maximum oxygen uptake in arms, leg, and combined arm-leg ergometry. J Appl Physiol 1975 May; 38(5): 774–9PubMedGoogle Scholar
  20. 20.
    Stenberg J, Astrand PO, Ekblom B, et al. Hemodynamic response to work with different muscle groups, sitting and supine. J Appl Physiol 1967 Jan; 22 (1): 61–70PubMedGoogle Scholar
  21. 21.
    Hermansen L, Saltin B. Oxygen uptake during maximal treadmill and bicycle exercise. J Appl Physiol 1969 Jan; 26(1): 31–7PubMedGoogle Scholar
  22. 22.
    Hermansen L, Ekblom B, Saltin B. Cardiac output during submaximal and maximal treadmill and bicycle exercise. J Appl Physiol 1970 Jul; 29 (1): 82–6PubMedGoogle Scholar
  23. 23.
    McArdle WD, Magel JR. Physical work capacity and maximum oxygen uptake in treadmill and bicycle exercise. Med Sci Sports 1970 Fall; 2 (3): 118–23PubMedGoogle Scholar
  24. 24.
    Faulkner JA, Roberts DE, Elk RL, et al. Cardiovascular responses to submaximum and maximum effort cycling and running. J Appl Physiol 1971 Apr; 30 (4): 457–61PubMedGoogle Scholar
  25. 25.
    Katch FI, McArdle WD, Pechar GS. Relationship of maximal leg force and leg composition to treadmill andbicycle ergometer maximum oxygen uptake. Med Sci Sports 1974 Spring; 6 (1): 38–43PubMedGoogle Scholar
  26. 26.
    Davis JA, Vodak P, Wilmore JH, et al. Anaerobic threshold and maximal aerobic power for three modes of exercise. J Appl Physiol 1976 Oct; 41 (4): 544–50PubMedGoogle Scholar
  27. 27.
    Hagberg JM, Giese MD, Schneider RB. Comparison of the three procedures for measuring VO2max in competitive cyclists. Eur J Appl Physiol Occup Physiol 1978 Jul 17; 39 (1): 47–52PubMedCrossRefGoogle Scholar
  28. 28.
    Matsui H, Kitamura K, Miyamura M. Oxygen uptake and blood flow of the lower limb in maximal treadmill and bicycle exercise. Eur J Appl Physiol Occup Physiol 1978 Dec 15; 40 (1): 57–62PubMedCrossRefGoogle Scholar
  29. 29.
    Miles DS, Critz JB, Knowlton RG. Cardiovascular, metabolic, and ventilatory responses of women to equivalent cycle ergometer and treadmill exercise. Med Sci Sports Exerc 1980 Spring; 12 (1): 14–9PubMedGoogle Scholar
  30. 30.
    Moreira-da-Costa M, Russo AK, Picarro IC, et al. Maximal oxygen uptake during exercise using trained or untrained muscles. Braz J Med Biol Res 1984; 17 (2): 197–202PubMedGoogle Scholar
  31. 31.
    Jacobs I, Sjodin B. Relationship of ergometer-specific VO2max and muscle enzymes to blood lactate during submaximal exercise. Br J Sports Med 1985 Jun; 19 (2): 77–80PubMedCrossRefGoogle Scholar
  32. 32.
    Moreira-da-Costa M, Russo AK, Picarro IC, et al. Oxygen consumption and ventilation during constant-load exercise in runners and cyclists. J Sports Med Phys Fitness 1989 Mar; 29 (1): 36–44Google Scholar
  33. 33.
    Green HJ, Sutton J, Young P, et al. Operation Everest II: muscle energetics during maximal exhaustive exercise. J Appl Physiol 1989 Jan; 66 (1): 142–50PubMedGoogle Scholar
  34. 34.
    Bouckaert J, Vrijens J, Pannier JL. Effect of specific test procedures on plasma lactate concentration and peak oxygen uptake in endurance athletes. J Sports Med Phys Fitness 1990 Mar; 30 (1): 13–8PubMedGoogle Scholar
  35. 35.
    Hill DW, Halcomb JN, Stevens EC. Oxygen uptake kinetics during severe intensity running and cycling. Eur J Appl Physiol 2003 Aug; 89 (6): 612–8PubMedCrossRefGoogle Scholar
  36. 36.
    Scott CB, Littlefield ND, Chason JD, et al. Differences in oxygen uptake but equivalent energy expenditure between a brief bout of cycling and running. Nutr Metab (Lond) 2006; 3: 1CrossRefGoogle Scholar
  37. 37.
    Bentley DJ, Newell J, Bishop D. Incremental exercise test design and analysis: implications for performance diagnostics in endurance athletes. Sports Med 2007; 37 (7): 575–86PubMedCrossRefGoogle Scholar
  38. 38.
    Midgley AW, Bentley DJ, Luttikholt H, et al. Challenging a dogma of exercise physiology: does an incremental exercise test for valid determination really need to last between 8-12 minutes? Sports Med 2008; 38 (6): 441–63PubMedCrossRefGoogle Scholar
  39. 39.
    Stromme SB, Ingjer F, Meen HD. Assessment of maximal aerobic power in specifically trained athletes. J Appl Physiol 1977 Jun; 42 (6): 833–7PubMedGoogle Scholar
  40. 40.
    Ricci J, Leger LA. VO2max of cyclists from treadmill, bicycle ergometer and velodrome tests. Eur J Appl PhysiolOccup Physiol 1983; 50 (2): 283–9PubMedCrossRefGoogle Scholar
  41. 41.
    Coyle EF, Coggan AR, Hopper MK, et al. Determinants of endurance in well-trained cyclists. J Appl Physiol 1988 Jun; 64 (6): 2622–30PubMedGoogle Scholar
  42. 42.
    Mazzeo RS, Marshall P. Influence of plasma catecholamines on the lactate threshold during graded exercise. J Appl Physiol 1989 Oct; 67 (4): 1319–22PubMedGoogle Scholar
  43. 43.
    Kravitz L, Robergs RA, Heyward VH, et al. Exercise mode and gender comparisons of energy expenditure at self-selected intensities. Med Sci Sports Exerc 1997 Aug; 29(8): 1028–35PubMedCrossRefGoogle Scholar
  44. 44.
    Zeni AI, Hoffman MD, Clifford PS. Energy expenditure with indoor exercise machines. JAMA 1996 May 8; 275(18): 1424–7PubMedCrossRefGoogle Scholar
  45. 45.
    Sedlock DA. Post-exercise energy expenditure after cycle ergometer and treadmill exercise. J Appl Sport Sci Res 1992; 6: 19–23Google Scholar
  46. 46.
    Coyle EF. Integration of the physiological factors determining endurance performance ability. Exerc Sport Sci Rev 1995; 23: 25–63PubMedCrossRefGoogle Scholar
  47. 47.
    Robergs RA, Ghiasvand F, Parker D. Biochemistry of exercise-induced metabolic acidosis. Am J Physiol Regul Integr Comp Physiol 2004 Sep; 287 (3): R502–16CrossRefGoogle Scholar
  48. 48.
    Loat CE, Rhodes EC. Relationship between the lactate and ventilatory thresholds during prolonged exercise. Sports Med 1993 Feb; 15 (2): 104–15PubMedCrossRefGoogle Scholar
  49. 49.
    Svedahl K, MacIntosh BR. Anaerobic threshold: the concept and methods of measurement. Can J Appl Physiol 2003 Apr; 28 (2): 299–323PubMedCrossRefGoogle Scholar
  50. 50.
    Hawley JA, Stepto NK. Adaptations to training in endurance cyclists: implications for performance. Sports Med 2001; 31 (7): 511–20PubMedCrossRefGoogle Scholar
  51. 51.
    Hassmen P. Perceptual and physiological responses to cycling and running in groups of trained and untrained subjects. Eur J Appl Physiol Occup Physiol 1990; 60 (6): 445–51PubMedCrossRefGoogle Scholar
  52. 52.
    Albrecht TL, Foster VL, Dickinson AL. Triathletes: exercise parameters measured during bicycle, swim bench,and treadmill testing [abstract]. Med Sci Sports Exerc 1986; 18: S86Google Scholar
  53. 53.
    Kohrt WM, Morgan DW, Bates B, et al. Physiological responses of triathletes to maximal swimming, cycling, and running. Med Sci Sports Exerc 1987 Feb; 19 (1): 51–5PubMedGoogle Scholar
  54. 54.
    O’Toole ML, Hiller DB, Crosby LO, et al. The ultra-endurance triathlete: a physiological profile. Med Sci Sports Exerc 1987 Feb; 19 (1): 45–50PubMedGoogle Scholar
  55. 55.
    O’Toole M, Hiller WDB, Douglas PS. Cardiovascular responses to prolonged cycling and running. Ann Sports Med 1987; 3: 124–30Google Scholar
  56. 56.
    Roalstad MS. Physiologic testing of the ultra-endurance triathlete. Med Sci Sports Exerc 1989 Oct; 21 (5 Suppl.): S200–4Google Scholar
  57. 57.
    Flynn MG, Costill DL, Kirwan JP, et al. Muscle fiber composition and respiratory capacity in triathletes. Int J Sports Med 1987 Dec; 8 (6): 383–6PubMedCrossRefGoogle Scholar
  58. 58.
    Kreider RB, Boone T, Thompson WR, et al. Cardiovascular and thermal responses of triathlon performance. Med Sci Sports Exerc 1988 Aug; 20 (4): 385–90PubMedCrossRefGoogle Scholar
  59. 59.
    Loftin M, Warren BL, Zingraf S, et al. Peak physiological function and performance of recreational triathletes. J Sports Med Phys Fitness 1988 Dec; 28 (4): 330–5PubMedGoogle Scholar
  60. 60.
    Dengel DR, Flynn MG, Costill DL, et al. Determinants of success during triathlon competition. Res Q Exerc Sport 1989 Sep; 60 (3): 234–8PubMedGoogle Scholar
  61. 61.
    Stein TP, Hoyt RW, Toole MO, et al. Protein and energy metabolism during prolonged exercise in trained athletes. Int J Sports Med 1989 Oct; 10 (5): 311–6PubMedCrossRefGoogle Scholar
  62. 62.
    Kohrt WM, O’Connor JS, Skinner JS. Longitudinal assessment of responses by triathletes to swimming, cycling, and running. Med Sci Sports Exerc 1989 Oct; 21 (5): 569–75PubMedGoogle Scholar
  63. 63.
    Millard-Stafford M, Sparling PB, Rosskopf LB, et al. Carbohydrate-electrolyte replacement during a simulated triathlon in the heat. Med Sci Sports Exerc 1990 Oct; 22(5): 621–8PubMedCrossRefGoogle Scholar
  64. 64.
    Rehrer NJ, Brouns F, Beckers EJ, et al. Gastric emptying with repeated drinking during running and bicycling. Int J Sports Med 1990 Jun; 11 (3): 238–43PubMedCrossRefGoogle Scholar
  65. 65.
    Butts NK, Henry BA, McLean D. Correlations between VO2max and performance times of recreational triathletes. J Sports Med Phys Fitness 1991 Sep; 31 (3): 339–44PubMedGoogle Scholar
  66. 66.
    Deitrick RW. Physiological responses of typical versus heavy weight triathletes to treadmill and bicycle exercise. J Sports Med Phys Fitness 1991 Sep; 31 (3): 367–75PubMedGoogle Scholar
  67. 67.
    Medelli J, Maingourd Y, Bouferrache B, et al. Maximal oxygen uptake and aerobic-anaerobic transition on treadmill and bicycle in triathletes. Jpn J Physiol 1993; 43(3): 347–60PubMedCrossRefGoogle Scholar
  68. 68.
    Sleivert GG, Wenger HA. Physiological predictors of short-course triathlon performance. Med Sci Sports Exerc 1993 Jul; 25 (7): 871–6PubMedCrossRefGoogle Scholar
  69. 69.
    Miura H, Ishiko T. Cardiorespiratory responses during a simulated triathlon. International council for health, physical education and recreation (ICHPER) 36th World Congress; 1993; Yokohama: 157–61Google Scholar
  70. 70.
    Murdoch SD, Bazzarre TL, Snider IP, et al. Differences in the effects of carbohydrate food form on endurance performance to exhaustion. Int J Sport Nutr 1993 Mar; 3 (1): 41–54PubMedGoogle Scholar
  71. 71.
    Miura H, Kitagawa K, Ishiko T, et al. Characteristics of VO2max and ventilatory threshold in triathletes. Jpn J Exerc Sports Physiol 1994; 1 (1): 99–106Google Scholar
  72. 72.
    Zhou S, Robson SJ, King MJ, et al. Correlations between short-course triathlon performance and physiological variables determined in laboratory cycle and treadmill tests. J Sports Med Phys Fitness 1997 Jun; 37 (2): 122–30PubMedGoogle Scholar
  73. 73.
    Roberts A, McElligott M. The relationship between strength and endurance in female triathletes. NSRC Scientific Report. Canberra (ACT): University of Canberra, 1995Google Scholar
  74. 74.
    Ruby B, Robergs R, Leadbetter G, et al. Cross-training between cycling and running in untrained females. J Sports Med Phys Fitness 1996 Dec; 36 (4): 246–54PubMedGoogle Scholar
  75. 75.
    Kerr CG, Trappe TA, Starling RD, et al. Hyperthermia during Olympic triathlon: influence of body heat storage during the swimming stage. Med Sci Sports Exerc 1998 Jan; 30 (1): 99–104PubMedCrossRefGoogle Scholar
  76. 76.
    Derman KD, Hawley JA, Noakes TD, et al. Fuel kinetics during intense running and cycling when fed carbohydrate. Eur J Appl Physiol Occup Physiol 1996; 74 (1-2): 36–43PubMedCrossRefGoogle Scholar
  77. 77.
    Miura H, Kitagawa K, Ishiko T. Economy during a simulated laboratory test triathlon is highly related to Olympic distance triathlon. Int J Sports Med 1997 May; 18 (4): 276–80PubMedCrossRefGoogle Scholar
  78. 78.
    Hue O, Le Gallais D, Chollet D, et al. The influence of prior cycling on biomechanical and cardiorespiratory response profiles during running in triathletes. Eur J Appl Physiol Occup Physiol 1998; 77 (1-2): 98–105PubMedCrossRefGoogle Scholar
  79. 79.
    Hue O, Le Gallais D, Boussana A, et al. Ventilatory responses during experimental cycle-run transition in triathletes. Med Sci Sports Exerc 1999 Oct; 31 (10): 1422–8PubMedCrossRefGoogle Scholar
  80. 80.
    Miura H, Kitagawa K, Ishiko T. Characteristic feature of oxygen cost at simulated laboratory triathlon test intrained triathletes. J Sports Med Phys Fitness 1999 Jun; 39(2): 101–6PubMedGoogle Scholar
  81. 81.
    Schabort EJ, Killian SC, St Clair Gibson A, et al. Prediction of triathlon race time from laboratory testing in national triathletes. Med Sci Sports Exerc 2000 Apr; 32 (4): 844–9PubMedCrossRefGoogle Scholar
  82. 82.
    Hue O, Le Gallais D, Boussana A, et al. Performance level and cardiopulmonary responses during a cycle-run trial. Int J Sports Med 2000 May; 21 (4): 250–5PubMedCrossRefGoogle Scholar
  83. 83.
    Toraa M, Friemel F. Fatigue of the respiratory muscles due to maximal exercise on 2 different ergometers. Can J Appl Physiol 2000 Apr; 25 (2): 87–101PubMedCrossRefGoogle Scholar
  84. 84.
    Hue O, Le Gallais D, Boussana A, et al. Catecholamine, blood lactate and ventilatory responses to multi-cycle-run blocks. Med Sci Sports Exerc 2000 Sep; 32 (9): 1582–6PubMedGoogle Scholar
  85. 85.
    Hue O, Le Gallais D, Prefaut C. Specific pulmonary responses during the cycle-run succession in triathletes. Scand J Med Sci Sports 2001 Dec; 11 (6): 355–61PubMedCrossRefGoogle Scholar
  86. 86.
    Hue O, Galy O, Le Gallais D, et al. Pulmonary responses during the cycle-run succession in elite and competitive triathletes. Can J Appl Physiol 2001 Dec; 26 (6): 559–73PubMedCrossRefGoogle Scholar
  87. 87.
    Vercruyssen F, Brisswalter J, Hausswirth C, et al. Influence of cycling cadence on subsequent running performance in triathletes. Med Sci Sports Exerc 2002 Mar; 34 (3): 530–6PubMedCrossRefGoogle Scholar
  88. 88.
    Basset F, Boulay MR. Treadmill and cycle ergometer tests are interchangeable to monitor triathletes annual training. J Sports Sci Med 2003; 2 (3): 110–6Google Scholar
  89. 89.
    Vercruyssen F, Suriano R, Bishop D, et al. Cadence selection affects metabolic responses during cycling and subsequent running time to fatigue. Br J Sports Med 2005 May; 39 (5): 267–72PubMedCrossRefGoogle Scholar
  90. 90.
    Schneider DA, Pollack J. Ventilatory threshold and maximal oxygen uptake during cycling and running in female triathletes. Int J Sports Med 1991 Aug; 12 (4): 379–83PubMedCrossRefGoogle Scholar
  91. 91.
    O’Toole ML, Douglas PS. Applied physiology of triathlon. Sports Med 1995 Apr; 19 (4): 251–67PubMedCrossRefGoogle Scholar
  92. 92.
    Miura H, Kitagawa K, Ishiko T. Characteristics of cardiorespiratory responses to the latter stage of a simulated triathlon. Jpn J Phys Fitness Sports Med 1994; 43: 381–8Google Scholar
  93. 93.
    De Vito G, Bernardi M, Sproviero E, et al. Decrease of endurance performance during Olympic triathlon. Int J Sports Med 1995 Jan; 16 (1): 24–8PubMedCrossRefGoogle Scholar
  94. 94.
    Billat VL, Mille-Hamard L, Petit B, et al. The role of cadence on the VO2 slow component in cycling andrunning in triathletes. Int J Sports Med 1999 Oct; 20 (7): 429–37PubMedCrossRefGoogle Scholar
  95. 95.
    Bernard T, Vercruyssen F, Grego F, et al. Effect of cycling cadence on subsequent 3 km running performance in well trained triathletes. Br J Sports Med 2003 Apr; 37 (2): 154-18; discussion 9PubMedCrossRefGoogle Scholar
  96. 96.
    Galy O, Hue O, Boussana A, et al. Effects of the order of running and cycling of similar intensity and duration on pulmonary diffusing capacity in triathletes. Eur J Appl Physiol 2003 Nov; 90 (5-6): 489–95PubMedCrossRefGoogle Scholar
  97. 97.
    Millet GP, Bentley DJ. The physiological responses to running after cycling in elite junior and senior triathletes. Int J Sports Med 2004 Apr; 25 (3): 191–7PubMedCrossRefGoogle Scholar
  98. 98.
    Galy O, Manetta J, Coste O, et al. Maximal oxygen uptake and power of lower limbs during a competitive season in triathletes. Scand J Med Sci Sports 2003 Jun; 13 (3): 185–93PubMedCrossRefGoogle Scholar
  99. 99.
    Bassett Jr DR, Howley ET. Limiting factors for maximum oxygen uptake and determinants of endurance performance. Med Sci Sports Exerc 2000 Jan; 32 (1): 70–84PubMedGoogle Scholar
  100. 100.
    Sloniger MA, Cureton KJ, Prior BM, et al. Lower extremity muscle activation during horizontal and uphill running. J Appl Physiol 1997 Dec; 83 (6): 2073–9PubMedGoogle Scholar
  101. 101.
    Bolognesi M. Ventilatory threshold and maximal oxygen uptake during cycling and running in duathletes. Med Sport 1997; 50: 209–16Google Scholar
  102. 102.
    Vleck VE, Bentley DJ, Millet GP, et al. Pacing during an elite Olympic distance triathlon: comparison between male and female competitors. J Sci Med Sport 2008; 11(4): 424–32PubMedCrossRefGoogle Scholar
  103. 103.
    Roecker K, Striegel H, Dickhuth HH. Heart-rate recommendations: transfer between running and cycling exercise? Int J Sports Med 2003 Apr; 24 (3): 173–8PubMedCrossRefGoogle Scholar
  104. 104.
    DiCarlo LJ, Sparling PB, Millard-Stafford ML, et al. Peak heart rates during maximal running and swimming: implications for exercise prescription. Int J Sports Med 1991 Jun; 12 (3): 309–12CrossRefGoogle Scholar
  105. 105.
    O’Toole ML, Douglas PS, Hiller WD. Use of heart rate monitors by endurance athletes: lessons from triathletes. J Sports Med Phys Fitness 1998 Sep; 38 (3): 181–7PubMedGoogle Scholar
  106. 106.
    Ray CA, Cureton KJ, Ouzts HG. Postural specificity of cardiovascular adaptations to exercise training. J Appl Physiol 1990 Dec; 69 (6): 2202–8PubMedGoogle Scholar
  107. 107.
    Kenny GP, Reardon FD, Marion A, et al. A comparative analysis of physiological responses at submaximal workloads during different laboratory simulations of field cycling. Eur J Appl Physiol Occup Physiol 1995; 71 (5): 409–15PubMedCrossRefGoogle Scholar
  108. 108.
    Gilman MB. The use of heart rate to monitor the intensity of endurance training. Sports Med 1996 Feb; 21 (2): 73–9PubMedCrossRefGoogle Scholar
  109. 109.
    Foster C, Lucia A. Running economy: the forgotten factor in elite performance. Sports Med 2007; 37 (4-5): 316–9PubMedCrossRefGoogle Scholar
  110. 110.
    Saunders PU, Pyne DB, Telford RD, et al. Factors affecting running economy in trained distance runners. Sports Med 2004; 34 (7): 465–85PubMedCrossRefGoogle Scholar
  111. 111.
    Hausswirth C, Bigard AX, Berthelot M, et al. Variability in energy cost of running at the end of a triathlon and a marathon. Int J Sports Med 1996 Nov; 17 (8): 572–9PubMedCrossRefGoogle Scholar
  112. 112.
    Hausswirth C, Bigard AX, Guezennec CY. Relationships between running mechanics and energy cost of running at the end of a triathlon and a marathon. Int J Sports Med 1997 Jul; 18 (5): 330–9PubMedCrossRefGoogle Scholar
  113. 113.
    Hausswirth C, Brisswalter J, Vallier JM, et al. Evolution of electromyographic signal, running economy, and perceived exertion during different prolonged exercises. Int J Sports Med 2000 Aug; 21 (6): 429–36PubMedCrossRefGoogle Scholar
  114. 114.
    Hausswirth C, Lehenaff D. Physiological demands of running during long distance runs and triathlons. Sports Med 2001; 31 (9): 679–89PubMedCrossRefGoogle Scholar
  115. 115.
    Guezennec CY, Vallier JM, Bigard AX, et al. Increase in energy cost of running at the end of a triathlon. Eur J Appl Physiol Occup Physiol 1996; 73 (5): 440–5PubMedCrossRefGoogle Scholar
  116. 116.
    Millet GP, Millet GY, Hofmann MD, et al. Alterations in running economy and mechanics after maximal cycling in triathletes: influence of performance level. Int J Sports Med 2000 Feb; 21 (2): 127–32PubMedCrossRefGoogle Scholar
  117. 117.
    Boone T, Kreider RB. Bicycle exercise before running: effect on performance. Ann Sports Med 1986; 3: 25–9Google Scholar
  118. 118.
    Millet GP, Vleck VE. Physiological and biomechanical adaptations to the cycle to run transition in Olympic triathlon: review and practical recommendations for training. Br J Sports Med 2000 Oct; 34 (5): 384–90PubMedCrossRefGoogle Scholar
  119. 119.
    Millet GP, Millet GY, Candau RB. Duration and seriousness of running mechanics alterations after maximal cycling in triathletes: influence of the performance level. J Sports Med Phys Fitness 2001 Jun; 41 (2): 147–53PubMedGoogle Scholar
  120. 120.
    Jones AM. The physiology of the world record holder for the women’s marathon. Int J Sports Sci Coaching 2006; 1 (2): 101–15CrossRefGoogle Scholar
  121. 121.
    Lucia A, Esteve-Lanao J, Olivan J, et al. Physiological characteristics of the best Eritrean runners-exceptional running economy. Appl Physiol Nutr Metab 2006 Oct; 31 (5): 530–40PubMedCrossRefGoogle Scholar
  122. 122.
    Lucia A, Olivan J, Bravo J, et al. The key to top-level endurance running performance: a unique example. Br J Sports Med 2008; 42 (3): 172–4PubMedCrossRefGoogle Scholar
  123. 123.
    Billat V, Lepretre PM, Heugas AM, et al. Training and bioenergetic characteristics in elite male and female Kenyan runners. Med Sci Sports Exerc 2003 Feb; 35 (2): 297–304; discussion 5-6PubMedCrossRefGoogle Scholar
  124. 124.
    Billat VL, Demarle A, Slawinski J, et al. Physical and training characteristics of top-class marathon runners. Med Sci Sports Exerc 2001 Dec; 33 (12): 2089–97PubMedCrossRefGoogle Scholar
  125. 125.
    Conley DL, Krahenbuhl GS, Burkett LN, et al. Following Steve Scott: physiological changes accompanying training. Phys Sportsmed 1984; 12: 103–6Google Scholar
  126. 126.
    Saltin B, Larsen H, Terrados N, et al. Aerobic exercise capacity at sea level and at altitude in Kenyan boys, junior and senior runners compared with Scandinavian runners. Scand J Med Sci Sports 1995 Aug; 5 (4): 209–21PubMedCrossRefGoogle Scholar
  127. 127.
    Saltin B, Kim CK, Terrados N, et al. Morphology, enzyme activities and buffer capacity in leg muscles of Kenyan and Scandinavian runners. Scand J Med Sci Sports 1995 Aug; 5 (4): 222–30PubMedCrossRefGoogle Scholar
  128. 128.
    Gaesser GA, Poole DC. The slow component of oxygen uptake kinetics in humans. Exerc Sport Sci Rev 1996; 24: 35–71PubMedCrossRefGoogle Scholar
  129. 129.
    Bijker KE, De Groot G, Hollander AP. Delta efficiencies of running and cycling. Med Sci Sports Exerc 2001 Sep; 33 (9): 1546–51PubMedCrossRefGoogle Scholar
  130. 130.
    Asmussen E, Bonde-Petersen F. Apparent efficiency and storage of elastic energy in human muscles during exercise. Acta Physiol Scand 1974 Dec; 92 (4): 537–45PubMedCrossRefGoogle Scholar
  131. 131.
    Zacks RM. The mechanical efficiencies of running and bicycling against a horizontal impeding force. Int Z Angew Physiol 1973 Jul 20; 31 (4): 249–58PubMedGoogle Scholar
  132. 132.
    Avela J, Kyrolainen H, Komi PV, et al. Reduced reflex sensitivity persists several days after long-lasting stretch-shortening cycle exercise. J Appl Physiol 1999 Apr; 86 (4): 1292–300PubMedGoogle Scholar
  133. 133.
    Farley CT, Gonzalez O. Leg stiffness and stride frequency in human running. J Biomech 1996 Feb; 29 (2): 181–6PubMedCrossRefGoogle Scholar
  134. 134.
    Kram R. Muscular force or work: what determines the metabolic energy cost of running? Exerc Sport Sci Rev 2000 Jul; 28 (3): 138–43PubMedGoogle Scholar
  135. 135.
    Kram R, Taylor CR. Energetics of running: a new perspective. Nature 1990 Jul 19; 346 (6281): 265–7PubMedCrossRefGoogle Scholar
  136. 136.
    Richardson RS, Harms CA, Grassi B, et al. Skeletal muscle: master or slave of the cardiovascular system? Med Sci Sports Exerc 2000 Jan; 32 (1): 89–93PubMedGoogle Scholar
  137. 137.
    di Prampero PE. Factors limiting maximal performance in humans. Eur J Appl Physiol 2003 Oct; 90 (3-4): 420–9PubMedCrossRefGoogle Scholar
  138. 138.
    Noakes TD. Maximal oxygen uptake: “classical” versus “contemporary” viewpoints: a rebuttal. Med Sci Sports Exerc 1998 Sep; 30 (9): 1381–98PubMedGoogle Scholar
  139. 139.
    Levine BD. VO2max: what do we know, and what do we still need to know? J Physiol 2008; 586: 25–34PubMedCrossRefGoogle Scholar
  140. 140.
    Prefaut C, Durand F, Mucci P, et al. Exercise-induced arterial hypoxaemia in athletes: a review. Sports Med 2000 Jul; 30 (1): 47–61PubMedCrossRefGoogle Scholar
  141. 141.
    Galy O, Le Gallais D, Hue O, et al. Is exercise-induced arterial hypoxemia in triathletes dependent on exercise modality? Int J Sports Med 2005 Nov; 26 (9): 719–26PubMedCrossRefGoogle Scholar
  142. 142.
    Powers SK, Lawler J, Dempsey JA, et al. Effects of incomplete pulmonary gas exchange on VO2 max. J Appl Physiol 1989 Jun; 66 (6): 2491–5PubMedGoogle Scholar
  143. 143.
    Green HJ, Carter S, Grant S, et al. Vascular volumes and hematology in male and female runners and cyclists. Eur J Appl Physiol Occup Physiol 1999 Feb; 79 (3): 244–50PubMedCrossRefGoogle Scholar
  144. 144.
    Galy O, Hue O, Boussana A, et al. Blood rheological responses to running and cycling: a potential effect on the arterial hypoxemia of highly trained athletes? Int J Sports Med 2005 Jan-Feb; 26 (1): 9–15PubMedCrossRefGoogle Scholar
  145. 145.
    Boussana A, Galy O, Hue O, et al. The effects of prior cycling and a successive run on respiratory muscle performance in triathletes. Int J Sports Med 2003 Jan; 24(1): 63–70PubMedCrossRefGoogle Scholar
  146. 146.
    Boussana A, Hue O, Matecki S, et al. The effect of cycling followed by running on respiratory muscle performance inelite and competition triathletes. Eur J Appl Physiol 2002 Aug; 87 (4-5): 441–7PubMedCrossRefGoogle Scholar
  147. 147.
    Boussana A, Matecki S, Galy O, et al. The effect of exercise modality on respiratory muscle performance in triathletes. Med Sci Sports Exerc 2001 Dec; 33 (12): 2036–43PubMedCrossRefGoogle Scholar
  148. 148.
    Hue O, Boussana A, Le Gallais D, et al. Pulmonary function during cycling and running in triathletes. J Sports Med Phys Fitness 2003 Mar; 43 (1): 44–50PubMedGoogle Scholar
  149. 149.
    Smith TB, Hopkins WG, Taylor NA. Respiratory responses of elite oarsmen, former oarsmen, and highly trained non-rowers during rowing, cycling and running. Eur J Appl Physiol Occup Physiol 1994; 69 (1): 44–9PubMedCrossRefGoogle Scholar
  150. 150.
    Gavin TP, Stager JM. The effect of exercise modality on exercise-induced hypoxemia. Respir Physiol 1999 May 3; 115 (3): 317–23PubMedCrossRefGoogle Scholar
  151. 151.
    Hopkins SR, Barker RC, Brutsaert TD, et al. Pulmonary gas exchange during exercise in women: effects of exercise type and work increment. J Appl Physiol 2000 Aug; 89 (2): 721–30PubMedGoogle Scholar
  152. 152.
    Hill NS, Jacoby C, Farber HW. Effect of an endurance triathlon on pulmonary function. Med Sci Sports Exerc 1991 Nov; 23 (11): 1260–4PubMedGoogle Scholar
  153. 153.
    Bonsignore MR, Morici G, Abate P, et al. Ventilation and entrainment of breathing during cycling and running in triathletes. Med Sci Sports Exerc 1998 Feb; 30 (2): 239–45PubMedCrossRefGoogle Scholar
  154. 154.
    Ekblom B. Effect of physical training on oxygen transport system in man. Acta Physiol Scand Suppl 1968; 328: 1–45PubMedGoogle Scholar
  155. 155.
    Saltin B, Blomqvist G, Mitchell JH, et al. Response to exercise after bed rest and after training. Circulation 1968 Nov; 38 (5 Suppl.): VII1–78Google Scholar
  156. 156.
    Delp MD, Laughlin MH. Regulation of skeletal muscle perfusion during exercise. Acta Physiol Scand 1998 Mar; 162 (3): 411–9PubMedCrossRefGoogle Scholar
  157. 157.
    Laaksonen MS, Kivela R, Kyrolainen H, et al. Effects of exhaustive stretch-shortening cycle exercise on muscleblood flow during exercise. Acta Physiol (Oxf) 2006 Apr; 186 (4): 261–70CrossRefGoogle Scholar
  158. 158.
    Rowland TW. The circulatory response to exercise: role of the peripheral pump. Int J Sports Med 2001 Nov; 22 (8): 558–65PubMedCrossRefGoogle Scholar
  159. 159.
    Sheriff DD. Muscle pump function during locomotion: mechanical coupling of stride frequency and muscle blood flow. Am J Physiol Heart Circ Physiol 2003 Jun; 284 (6): H2185–91Google Scholar
  160. 160.
    Noakes TD, St Clair Gibson A. Logical limitations to the “catastrophe” models of fatigue during exercise in humans. Br J Sports Med 2004 Oct; 38 (5): 648–9PubMedCrossRefGoogle Scholar
  161. 161.
    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 Dec; 38 (6): 797–806PubMedCrossRefGoogle Scholar
  162. 162.
    Holloszy JO, Coyle EF. Adaptations of skeletal muscle to endurance exercise and their metabolic consequences. J Appl Physiol 1984 Apr; 56 (4): 831–8PubMedGoogle Scholar
  163. 163.
    Coyle EF, Feltner ME, Kautz SA, et al. Physiological and biomechanical factors associated with elite endurance cycling performance. Med Sci Sports Exerc 1991 Jan; 23 (1): 93–107PubMedGoogle Scholar
  164. 164.
    Weston AR, Myburgh KH, Lindsay FH, et al. Skeletal muscle buffering capacity and endurance performance after high-intensity interval training by well-trained cyclists. Eur J Appl Physiol Occup Physiol 1997; 75 (1): 7–13PubMedCrossRefGoogle Scholar
  165. 165.
    Green HJ, Patla AE. Maximal aerobic power: neuromuscular and metabolic considerations. Med Sci Sports Exerc 1992 Jan; 24 (1): 38–46PubMedGoogle Scholar
  166. 166.
    Aunola S, Marniemi J, Alanen E, et al. Muscle metabolic profile and oxygen transport capacity as determinants ofaerobic and anaerobic thresholds. Eur J Appl Physiol Occup Physiol 1988; 57 (6): 726–34PubMedCrossRefGoogle Scholar
  167. 167.
    Ivy JL, Costill DL, Maxwell BD. Skeletal muscle determinants of maximum aerobic power in man. Eur J Appl Physiol Occup Physiol 1980; 44 (1): 1–8PubMedCrossRefGoogle Scholar
  168. 168.
    Coyle EF, Sidossis LS, Horowitz JF, et al. Cycling efficiency is related to the percentage of type I muscle fibers. Med Sci Sports Exerc 1992 Jul; 24 (7): 782–8PubMedGoogle Scholar
  169. 169.
    Horowitz JF, Sidossis LS, Coyle EF. High efficiency of type I muscle fibers improves performance. Int J Sports Med 1994 Apr; 15 (3): 152–7PubMedCrossRefGoogle Scholar
  170. 170.
    Parkhouse WS, McKenzie DC, Hochachka PW, et al. Buffering capacity of deproteinized human vastus lateralis muscle. J Appl Physiol 1985 Jan; 58 (1): 14–7PubMedGoogle Scholar
  171. 171.
    Bijker KE, de Groot G, Hollander AP. Differences in leg muscle activity during running and cycling in humans. Eur J Appl Physiol 2002 Oct; 87 (6): 556–61PubMedCrossRefGoogle Scholar
  172. 172.
    Marcinik EJ, Potts J, Schlabach G, et al. Effects of strength training on lactate threshold and endurance performance. Med Sci Sports Exerc 1991 Jun; 23 (6): 739–43PubMedGoogle Scholar
  173. 173.
    Chapman AR, Vicenzino B, Blanch P, et al. Does cycling effect motor coordination of the leg during running in elite triathletes? J Sci Med Sport 2008; 11 (4): 371–80PubMedCrossRefGoogle Scholar
  174. 174.
    Borg G, Van den Burg M, Hassmen P. Relationships between perceived exertion, HR and HLa in cycling, running and walking. Scand J Sports Sci 1987; 9: 69–77Google Scholar
  175. 175.
    Marsh AP, Martin PE. Effect of cycling experience, aerobic power, and power output on preferred and most economical cycling cadences. Med Sci Sports Exerc 1997 Sep; 29 (9): 1225–32PubMedCrossRefGoogle Scholar
  176. 176.
    Patterson RP, Moreno MI. Bicycle pedalling forces as a function of pedalling rate and power output. Med Sci Sports Exerc 1990 Aug; 22 (4): 512–6PubMedGoogle Scholar
  177. 177.
    Takaishi T, Yasuda Y, Ono T, et al. Optimal pedaling rate estimated from neuromuscular fatigue for cyclists. Med Sci Sports Exerc 1996 Dec; 28 (12): 1492–7PubMedCrossRefGoogle Scholar
  178. 178.
    Lucia A, Hoyos J, Chicharro JL. Preferred pedalling cadence in professional cycling. Med Sci Sports Exerc 2001 Aug; 33 (8): 1361–6PubMedCrossRefGoogle Scholar
  179. 179.
    Marsh AP, Martin PE, Foley KO. Effect of cadence, cycling experience, and aerobic power on delta efficiency during cycling. Med Sci Sports Exerc 2000 Sep; 32 (9): 1630–4PubMedGoogle Scholar
  180. 180.
    Marsh AP, Martin PE. The relationship between cadence and lower extremity EMG in cyclists and noncyclists. Med Sci Sports Exerc 1995 Feb; 27 (2): 217–25PubMedGoogle Scholar
  181. 181.
    Lepers R, Hausswirth C, Maffiuletti N, et al. Evidence of neuromuscular fatigue after prolonged cycling exercise. Med Sci Sports Exerc 2000 Nov; 32 (11): 1880–6PubMedCrossRefGoogle Scholar
  182. 182.
    Vercruyssen F, Hausswirth C, Smith D, et al. Effect of exercise duration on optimal pedaling rate choice in triathletes. Can J Appl Physiol 2001 Feb; 26 (1): 44–54PubMedGoogle Scholar
  183. 183.
    Brisswalter J, Hausswirth C, Smith D, et al. Energetically optimal cadence vs. freely-chosen cadence during cycling:effect of exercise duration. Int J Sports Med 2000 Jan; 21 (1): 60–4PubMedCrossRefGoogle Scholar
  184. 184.
    Gottschall JS, Palmer BM. The acute effects of prior cycling cadence on running performance and kinematics. Med Sci Sports Exerc 2002 Sep; 34 (9): 1518–22PubMedCrossRefGoogle Scholar
  185. 185.
    Bentley DJ, Millet GP, Vleck VE, et al. Specific aspects of contemporary triathlon: implications for physiological analysis and performance. Sports Med 2002; 32 (6): 345–59PubMedCrossRefGoogle Scholar
  186. 186.
    Bernard T, Vercruyssen F, Mazure C, et al. Constant versus variable-intensity during cycling: effects on subsequent running performance. Eur J Appl Physiol 2007 Jan; 99 (2): 103–11PubMedCrossRefGoogle Scholar
  187. 187.
    Vleck VE, Burgi A, Bentley DJ. The consequences of swim, cycle, and run performance on overall result in eliteolympic distance triathlon. Int J Sports Med 2006 Jan; 27 (1): 43–8PubMedCrossRefGoogle Scholar
  188. 188.
    Millet GY, Lepers R. Alterations of neuromuscular function after prolonged running, cycling and skiing exercises. Sports Med 2004; 34 (2): 105–16PubMedCrossRefGoogle Scholar
  189. 189.
    Millet GY, Lepers R, Maffiuletti NA, et al. Alterations of neuromuscular function after an ultra marathon. J Appl Physiol 2002 Feb; 92 (2): 486–92PubMedGoogle Scholar
  190. 190.
    Millet GY, Martin V, Lattier G, et al. Mechanisms contributing to knee extensor strength loss after prolonged running exercise. J Appl Physiol 2003 Jan; 94 (1): 193–8PubMedGoogle Scholar
  191. 191.
    Lepers R, Maffiuletti NA, Rochette L, et al. Neuromuscular fatigue during a long-duration cycling exercise. J Appl Physiol 2002 Apr; 92 (4): 1487–93PubMedGoogle Scholar
  192. 192.
    Lepers R, Millet GY, Maffiuletti NA. Effect of cycling cadence on contractile and neural properties of knee extensors. Med Sci Sports Exerc 2001 Nov; 33 (11): 1882–8PubMedCrossRefGoogle Scholar
  193. 193.
    Racinais S, Girard O, Micallef JP, et al. Failed excitability of spinal motoneurons induced by prolonged running exercise. J Neurophysiol 2007 Jan; 97 (1): 596–603PubMedCrossRefGoogle Scholar
  194. 194.
    Millet GY, Millet GP, Lattier G, et al. Alteration of neuromuscular function after a prolonged road cycling race. Int J Sports Med 2003 Apr; 24 (3): 190–4PubMedCrossRefGoogle Scholar
  195. 195.
    Bentley DJ, Smith PA, Davie AJ, et al. Muscle activation of the knee extensors following high intensity endurance exercise in cyclists. Eur J Appl Physiol 2000 Mar; 81 (4): 297–302PubMedCrossRefGoogle Scholar
  196. 196.
    Takaishi T, Yasuda Y, Moritani T. Neuromuscular fatigue during prolonged pedalling exercise at different pedalling rates. Eur J Appl Physiol Occup Physiol 1994; 69 (2): 154–8PubMedCrossRefGoogle Scholar

Copyright information

© Springer International Publishing AG 2009

Authors and Affiliations

  • Gregoire P. Millet
    • 1
  • V. E. Vleck
    • 2
  • D. J. Bentley
    • 3
  1. 1.ISSEPUniversity of LausanneLausanneSwitzerland
  2. 2.School of BiosciencesUniversity of WestminsterLondonUK
  3. 3.Health and Exercise, School of Medical ScienceUniversity of New South WalesSydneyAustralia

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