Review
What makes an endurance athlete world-class? Not simply a physiological conundrum

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Abstract

Inter-individual variation in endurance performance capacity is a characteristic, not only of the general population, but also in trained athletes. The ability of sport scientists to predict which athletes amongst an elite group will become world-class is limited. We do not fully understand the interactions between biological factors, training, recovery and competitive performance. Assessment methods and interpretation of results do not take into account the facts that most research is not done on elite athletes and performances of world-class endurance athletes cannot be attributed to aerobic capacity alone. Many lines of evidence suggest that there is a limit to adaptation in aerobic capacity. Recent advances in molecular biology and genetics should be harnessed by exercise biologists in conjunction with previously used physiological, histological and biochemical techniques to study elite athletes and their responses to different training and recovery regimens. Technological advances should be harnessed to study world-class athletes to determine optimal training and competition strategies. In summary, it is likely that multiple factors are essential contributors to world-class endurance performance and that it is only by using a multidisciplinary approach that we will come closer to solving the conundrum: ‘What makes an endurance athlete world class?’

Introduction

In humans endurance performance capacity is characterized by a large degree of inter-individual variation in the general population (Bouchard et al., 1993) and even in well-trained, athletic individuals. This capacity was historically, typically assessed by exercise scientists using physiology, biochemistry and histology techniques (see Fig. 1). The most frequently used tests included determination of the whole body maximal and sub-maximal oxygen consumption and the lactate turnpoint (Costill, 1967), oxidative enzyme activities and the percentage of slow twitch fibers (Booth and Narahara, 1975, Bylund et al., 1977). All these parameters correlate well with endurance performance in events ranging from 3000 m to various ultramarathon distances in heterogeneous groups with elite endurance athletes clearly different from recreational runners (Larsen, 2003). But when the conundrum is posed: ‘What makes an endurance athlete world-class?’ one must ask whether the above-mentioned methods are sufficient for assessing and distinguishing between elite- and truly world-class athletes. The first point that this review will make is that the above measures are not effective as performance predictors amongst relatively homogenous groups of well-trained elite or even sub-elite athletes. Having ascertained this, new questions can be raised, e.g. ‘Are newer assessment methods more useful? Do other biological factors contribute to endurance performance?’

The second major objective is to review the literature that could help us to answer the above questions, focusing mainly on the past decade, and to update the schema presented in Fig. 1, appropriately. In particular, it will become clear that exercise scientists in the new millennium should at least understand, if not integrate into their research programs, contributions made in the fields of genetics and molecular biology. The third major objective is to provide a more integrated schema that reflects the multidisciplinary approach that I propose exercise scientists should take over the next few years. Fig. 1 also presents scientifically based training as a necessary component of elite endurance performance. However, are current training methods of elite athletes really scientifically based, or are scientific studies based on what elite athletes are already doing in training? The concepts in the central portion of the schema presented in Fig. 1 are certainly not new, but this review will argue, on the one hand, that many questions remain to be adequately answered and on the other hand, that new information needs to be added for us to understand what optimal training leading to truly world-class performance actually is. Therefore, as the review progresses, the framework will be updated to reflect the contributions of other branches of biology and new focuses of exercise science, and to reflect the dual necessities of studying elite athletes and using the most integrative, multidisciplinary approach possible in the future.

Although studies on endurance cyclists, tri-athletes and cross-country skiers have contributed substantially to our understanding of ‘the endurance athlete’, I will focus, where possible, particularly on studies on runners. The dominance of the Kenyan endurance runners in world competition has returned time and again to the stage of the popular media. Equally intriguing are the sheer numbers of African distance runners who at times have beaten, but more frequently follow closely in the Kenyans’ footsteps. This article will mention, where appropriate, those comparative studies that have, albeit in a fairly limited way, explored a physiological and/or biochemical basis for these phenomena. It is pertinent to address the fact that most studies in the scientific literature are on sub-elite, rather than elite or world-class athletes, and that this should be taken into account when interpreting the literature (cf. Larsen, 2003).

Section snippets

The in vivo physiology of endurance performance

Hill and Lupton (1923) suggested that Vo2max was higher in endurance-trained athletes when compared to less trained controls. Since this early study, many other studies over the years have concurred that, in general, Vo2max is a good predictor of endurance running performance (e.g. Costill, 1967, Foster et al., 1978, Sjödin and Svedenhag, 1985). But, this is not the case in groups of elite or sub-elite athletes with more homogeneous race performances (Conley and Krahenbuhl, 1980). It was also

Implications of the schematic framework

The central framework in Fig. 1, represents not only multiple important factors crucial for elite athletic performance, but also an analogy to illustrate their relative importance. The schema is analogous to a series of three-legged stools and this has two implications: firstly, if any one leg is absent, the stool will topple; secondly, if any one leg is unstable, the stool will collapse if placed under sufficient pressure. The three disciplines represented together as components of exercise

Factors related to endurance performance that are assessed in vitro

As mentioned above and indicated in Fig. 1, endurance performance is reliant on at least three components: physiology, biochemistry and histology. The ‘boundaries’ between physiology and biochemistry were bridged by exercise scientists many years ago (e.g. Foster et al., 1978), but for the purposes of this review I have discussed whole body assessments and plasma analyses of blood samples taken during exercise under physiology and will now discuss other assessments that are made on the basis of

The adapted framework

Fig. 1 illustrated at least three branches of scientific endeavor that were typically used to investigate scientific aspects of athletic performance. However, the mushrooming of research in genetics and other branches of biological science over the past decade includes many studies related to exercise and truly emphasizes its biological nature. To accommodate this change, Fig. 2 now represents histology as a branch of physiology and combines biochemistry and other branches of molecular biology

Molecular biology, genetics and endurance performance

In the previous section, I discussed adaptive responses to endurance training at the level of protein expression, organelle and muscle cell size focusing primarily on observations in the muscle of endurance trained athletes. Some of those data have been available for decades, so the purpose was to highlight new additions, new interpretations or neglected aspects. However, as stated in a recent review by Hoppeler and Flück (2002), we still know relatively little about the molecular signals

Factors outside of biology that influence world-class endurance performance

The biological factors discussed above are clearly permissive with respect to adaptations favorable for endurance performance. However, this review would not be complete without mentioning the possibility that factors outside of the biological may actually underpin world-class athletic performance.

The multidisciplinary and integrative framework for future research

Collins (2002) recently proposed that it is only by using a multidisciplinary and interdisciplinary approach that life scientists can cope with the current rapid changes in biology, and I propose that this is also true for exercise science. This review led to the development of the schema in Fig. 2 that presents the branches of biology as overlapping, inter-related disciplines. Exercise science is presented as a multidisciplinary research endeavor encompassing the various inter-related

References (141)

  • L.V. Billat

    Interval training for performance: a scientific and empirical practice. Special recommendations for middle- and long-distance running. Part I: aerobic interval training

    Sports Med.

    (2001)
  • V. Billat et al.

    Reproducibility of running time to exhaustion at Vo2max in sub-elite runners

    Med. Sci. Sports Exerc.

    (1994)
  • A. Bonen

    Lactate transporters (MCT proteins) in heart and skeletal muscles

    Med. Sci. Sports Exerc.

    (2000)
  • F.W. Booth et al.

    Vastus lateralis cytochrome oxidase activity and its relationship to maximal oxygen consumption in man

    Pflügers Arch.

    (1975)
  • C. Bouchard et al.

    Genetics of aerobic and anaerobic performances

    Exerc. Sport Sci. Rev.

    (1993)
  • L.J. Brandon

    Physiological factors associated with middle distance running performance

    Sports Med.

    (1995)
  • L.J. Brandon et al.

    Influence of metabolic, mechanical and physique variables on middle distance running

    J. Sports Med. Phys. Fitness

    (1992)
  • G.A. Brooks

    Lactate shuttle – between but not within cells?

    J. Physiol.

    (2002)
  • L.M. Burke et al.

    Muscle glycogen storage after prolonged exercise: effect of the glycemic index of carbohydrate feedings

    J. Appl. Physiol.

    (1993)
  • A.C. Bylund et al.

    Physical training in man-skeletal muscle metabolism in relation to muscle morphology and running ability

    Eur. J. Appl. Physiol.

    (1977)
  • R.B. Child et al.

    Effects of a training taper on tissue damage indices, serum antioxidant capacity and half-marathon running performance

    Int. J. Sports Med.

    (2000)
  • P. Coetzer et al.

    Superior fatigue resistance of elite black South African distance runners

    J. Appl. Physiol.

    (1993)
  • Coggan, A.R., Williams, B.D., 1995. Metabolic adaptations to endurance training. Substrate metabolism during exercise....
  • J.P. Collins

    May you live in interesting times: using multidisciplinary and interdisciplinary programs to cope with change in the life sciences

    Bioscience

    (2002)
  • D.L. Conley et al.

    Running economy and distance running performance of highly trained athletes

    Med. Sci. Sports Exerc.

    (1980)
  • K.E. Conley et al.

    Limits to sustainable muscle performance: interaction between glycolysis and oxidative phosphorylation

    J. Exp. Biol.

    (2001)
  • D.A. Connolly et al.

    Treatment and prevention of delayed onset muscle soreness

    J. Strength Cond. Res.

    (2003)
  • D.L. Costill

    The relationship between selected physiological variables and distance running performance

    J. Sports Med. Phys. Fitness

    (1967)
  • D.L. Costill et al.

    Muscle fiber composition and enzyme activities of elite distance runners

    Med. Sci. Sports

    (1976)
  • D.L. Costill et al.

    Fractional utilization of the aerobic capacity during distance running

    Med. Sci. Sports

    (1973)
  • D.L. Costill et al.

    Effects of repeated days of intensified training on muscle glycogen and swimming performance

    Med. Sci. Sports Exerc.

    (1988)
  • E.F. Coyle et al.

    Determinants of endurance in well-trained cyclists

    J. Appl. Physiol.

    (1988)
  • B. Dawson et al.

    Effect of vitamin C and E supplementation on biochemical and ultrastructural indices of muscle damage after a 21 km run

    Int. J. Sports Med.

    (2002)
  • W. Derman et al.

    The ‘worn-out athlete’: a clinical approach to chronic fatigue in athletes

    J. Sports Sci.

    (1997)
  • F.T. Dionne et al.

    Mitochondrial DNA sequence polymorphism, Vo2max, and response to endurance training

    Med. Sci. Sports Exerc.

    (1991)
  • B. Essen-Gustavsson et al.

    Enzyme levels in pools of microdissected human muscle fibres of identified type. Adaptive response to exercise

    Acta Physiol. Scand.

    (1984)
  • P.A. Farrell et al.

    Plasma lactate accumulation and distance running performance

    Med. Sci. Sports

    (1979)
  • R.A. Fielding et al.

    What are the dietary protein requirements of physically active individuals? New evidence on the effects of exercise on protein utilization during post-exercise recovery

    Nutr. Clin. Care

    (2002)
  • C. Foster et al.

    Skeletal muscle enzyme activity and Vo2max in relation to distance running performance

    Eur. J. Appl. Physiol.

    (1978)
  • C. Foster et al.

    Effect of pacing strategy on cycle time trial performance

    Med. Sci. Sports Exerc.

    (1993)
  • A.C. Fry et al.

    Resistance exercise overtraining and overreaching. Neuroendocrine responses

    Sports Med.

    (1997)
  • A.S. Gibson et al.

    Exercise-induced mitochondrial dysfunction in an elite athlete

    Clin. J. Sports Med.

    (1998)
  • S. Grant et al.

    The relationship between 3 km running performance and selected physiological variables

    J. Sports Sci.

    (1997)
  • T. Gustafsson et al.

    Exercise-induced expression of angiogenesis-related transcription and growth factors in human skeletal muscle

    Am. J. Physiol.

    (1999)
  • S.L. Halson et al.

    Time course of performance changes and fatigue markers during intensified training in trained cyclists

    J. Appl. Physiol.

    (2002)
  • S.L. Halson et al.

    Immunological responses to overreaching in cyclists

    Med. Sci. Sports Exerc.

    (2003)
  • M.P. Harber et al.

    Myosin heavy chain composition of single muscle fibers in male distance runners

    Int. J. Sports Med.

    (2002)
  • J.A. Hawley

    Adaptations of skeletal muscle to prolonged, intense endurance training

    Clin. Exp. Pharmacol. Physiol.

    (2002)
  • J.A. Hawley et al.

    High intensity training in previously well trained endurance cyclists

    J. Sports Sci.

    (1997)
  • R.T. Hepple et al.

    Resistance and aerobic training in older men: effects on Vo2peak and the capillary supply to skeletal muscle

    J. Appl. Physiol.

    (1997)
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