REVIEW ARTICLE

Sports Med 2001; 31 (5): 325-337 0112-1642/01/0005-0325/$22.00/0 Adis International Limited. All rights reserved.

Physiology of Professional Road Cycling

Alejandro Luca,1,2 Jess Hoyos2,3 and Jos L. Chicharro2,4
1 2 3 4 Department of Anatomy and Physiology, European University of Madrid, Madrid, Spain Exercise Physiology Research Unit, Complutense University of Madrid, Madrid, Spain iBanesto.com Cycling Team, Banesto Sport Association, Madrid, Spain Department of Nursery, School of Medicine, Complutense University of Madrid, Madrid, Spain

Contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. Main Characteristics of the Sport . . . . . . . . . . . . . . . . . . . . . . . 1.1 Racing Calendar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.2 Tour Races . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.2.1 Flat Stages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.2.2 Time Trials . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.2.3 High Mountain Ascents . . . . . . . . . . . . . . . . . . . . . . . 2. Anthropometric Variables . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. Physiological Variables . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.1 Maximal Power Output and Maximal Oxygen Uptake . . . . . . . . 3.2 Submaximal Variables . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.2.1 Anaerobic Threshold . . . . . . . . . . . . . . . . . . . . . . . . 3.2.2 Isocapnic Buffering and Hypocapnic Hyperventilation Phases 3.2.3 Oxygen Uptake Kinetics during Submaximal Exercise . . . . . 4. Cardiovascular Responses and Adaptations . . . . . . . . . . . . . . . . . 4.1 Heart Rate (HR) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.1.1 The HR Response to Incremental Exercise: the Conconi Test . 4.1.2 HR as an Indicator of Exercise Intensity . . . . . . . . . . . . . . 4.2 Cardiovascular Adaptations . . . . . . . . . . . . . . . . . . . . . . . 4.2.1 Cardiac Adaptations . . . . . . . . . . . . . . . . . . . . . . . . 4.2.2 Vascular Adaptations . . . . . . . . . . . . . . . . . . . . . . . . 5. Pulmonary Responses and Adaptations . . . . . . . . . . . . . . . . . . . 6. Physiological Adaptations during a Sports Season . . . . . . . . . . . . . 6.1 Maximal Variables . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.2 Submaximal Variables: Metabolic and Neuromuscular Adaptations 7. Endocrine System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. Haematological Variables and Blood Doping . . . . . . . . . . . . . . . . 9. Nutritional Habits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9.1 Tour Races . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9.2 Training . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10.Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 325 326 326 326 326 327 327 328 328 328 328 328 329 329 330 330 330 330 331 331 332 332 332 333 333 333 334 334 334 335 335

Abstract

Professional road cycling is an extreme endurance sport. Approximately 30 000 to 35 000km are cycled each year in training and competition and some races,

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such as the Tour de France last 21 days (~100 hours of competition) during which professional cyclists (PC) must cover >3500km. In some phases of such a demanding sport, on the other hand, exercise intensity is surprisingly high, since PC must complete prolonged periods of exercise (i.e. time trials, high mountain . ascents) at high percentages (~90%) of maximal oxygen uptake (V O2max) [above the anaerobic threshold (AT)]. Although numerous studies have analysed the physiological responses of elite, amateur level road cyclists during the last 2 decades, their findings might not be directly extrapolated to professional cycling. Several studies have recently shown that PC exhibit some remarkable physiological responses and adaptations such as: an efficient respiratory system (i.e. lack of tachypnoeic shift at high exercise intensities); a considerable reliance on fat metabolism even at high power outputs; or several neuromuscular adaptations (i.e. a great resistance to fatigue of slow motor units). This article extensively reviews the different responses and adaptations (cardiopulmonary system, metabolism, neuromuscular factors or endocrine system) to this sport. A special emphasis is placed on the evaluation of performance both in the laboratory (i.e. the controversial Conconi test, distinction between climbing and time trial ability, etc.) and during actual competitions such as the Tour de France.

1. Main Characteristics of the Sport

1.1 Racing Calendar

flat, long parcours (usually ridden at high speeds inside a large group of riders), individual time trials (TT) [40 to 60km on an overall level terrain] and uphill cycling (high mountain passes).[1-3]
1.2.1 Flat Stages

Approximately 30 000 to 35 000km are cycled each year by professional cyclists (PC) in both training and competition. The typical racing season of a PC starts in the late winter (mid-February) and finishes at the end of summer/beginning of fall. It comprises a total of 90 to 100 competition days, including: (i) numerous 1-day races (i.e. Classics of ~250km); (ii) several 1-week tour races (i.e. 4 to 5 consecutive daily stages of 150 to 200km and an individual time trial); and (iii) one or two 3-week tour races (i.e. Giro dItalia, Tour de France and Vuelta a Espaa). The latter include 21 daily stages (~200km or 4 to 5 hours per stage) with only 1 to 2 days of rest, that is, 90 to 100 hours of competition during which cyclists must cover 3500 to 4000km.
1.2 Tour Races

Professional road cycling is a complex sport in which many uncontrollable variables (weather conditions, altitude, wind direction, team tactics, etc.) can affect performance. Nevertheless, we can generalise that 3-week tour races such as the Tour de France include 3 main competition requirements:
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Every tour race includes seven or more flat stages of ~200km (4 to 5 hours) [table I]. In such long, flat stages, PC ride most of the time inside a large group of 150 to 200 fellow competitors which considerably reduces the primary force to be overcome in this type of terrain that is, air resistance. As a result, the energy requirement of cycling can be decreased by as much as 40%,[8] which makes overall exercise intensity low-to-moderate. Indeed, the percentage of the total stage time spent at an intensity below 70%, between 70 and 90%, and above 90% . of maximal oxygen uptake (VO2max), averages approximately 70, 25 and 5%, respectively.[2] Cycling cadence, on the other hand, averages ~90 rpm (data collected during 1999 in the Giro dItalia, Tour de France and Vuelta a Espaa).[4] A great mastery of technical skills (drafting or the ability to avoid crashes) appears most important in these types of stages, which most PC are able to finish in the same time and which usually do not determine the final outcome of tour races.[1] Nevertheless, the high average velocities (~45 km/h)
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Table I. Main characteristics of flat and high mountain stages, and time trials for professional cyclists in tour races[1-7] FS Distance (km) Exercise time (h) Mean exercise intensity Prevailing metabolism Mean velocity (km/h) Cadence (rpm) Cycling position Main requirements Specific concerns Estimated power output ~200 4-5 Low-to-moderate Aerobic (fat) ~45 ~90 Traditional (sitting) Technical Crashes, muscle damage <250W HMS ~200 5-6 Moderate-to-high (high during ascents) Aerobic (fat and CHO) and Aerobic/anaerobic (CHO) during ascents ~20 (during ascents) ~70 Alternating (sitting and standing) Physiological Moderate hypoxia 6 or more W/kg (during ascents) TT 40-60 (overall flat) ~1 High Aerobic (CHO) and anaerobic ~50 (TT specialists) ~90 Aerodynamic (triathlon bars) Physiological and aerodynamics Aerodynamics 350W (400W in TT specialists)

CHO = carbohydrate; FS = flat stages; HMS = high mountain stages; TT = time trials.

at which PC are able to cover these stages require that they push high gears (i.e. 53 12-11) during long periods. This inevitably results in some degree of muscle damage.[1] Previous research has indeed reported increased levels of muscle damage markers during cycling tour races.[9] This phenomenon might have a negative impact on performance during the second part of 3-week races, during which accumulated muscle fatigue may considerably limit performance in the most crucial phases of competition, that is, time trials (TT) and high mountain passes.
1.2.2 Time Trials

that the mean absolute power output of an average PC during long TT averages ~350W,[6] although TT specialists probably generate much higher power outputs (>400W).[3] Indeed, a mathematical approach recently allowed Bassett and co-workers[7] to estimate that the mean power outputs required to break the 1-hour world records in a velodrome during the last 7 years (53.0 to 56.4km) ranged between 427 and 460W. Padilla et al.[10] recently estimated that the mean power output corresponding to one of the 1994 1-hour records averaged as much as 509.5W.
1.2.3 High Mountain Ascents

Tour races usually include 3 TT which are held along overall flat terrains: a short, opening TT of 5 to 10km and 2 long TT (40 to 60km) [table I]. This phase of the competition is usually crucial in the final outcome of tour races. Air resistance is the main force that the cyclist encounters during TT and thus aerodynamic factors (cyclists riding posture, size of frontal wheels, etc.) play a major role.[5] Those PC who seek top performances (average velocity of ~50 km/h) must tolerate high constant workloads (close to the anaerobic threshold [AT] or ~90% . VO2max) during the entire TT (~60 minutes).[1] During these events, the percentage of total exercise time spent at an exercise intensity above 90% of . VO2max averages ~50%.[1,2] The best time trialists must push extremely demanding gears (i.e. 54-55 12-11) at very high rates (average of ~90 rpm) during long periods. Some authors have estimated
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Some group stages of ~200km (the so-called high mountain stages) include 3 to 5 mountain passes of 5 to 10% mean gradient and thus require cycling uphill during several periods of 30 to 60 minutes over a total time of 5 to 6 hours (table I). When climbing at low speeds [~20 km/h in hors category mountain passes (most difficult mountain passes according to tour race organisers)], the cyclist must mainly overcome the force of gravity.[11] Because of its effects on gravity-induced resistance, body mass has a major influence on climbing performance.[11] A high power output : body mass ratio at maximal or near-to-maximal intensities (6 or more W/kg) is thus a necessary prerequisite for PC.[3,5] In addition, rolling resistance resulting from the interaction between the bicycle tires and the road surface considerably increases at these lower riding speeds and when riding on rough road surfaces such as
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those of most mountain cols.[5] To overcome the aforementioned forces, cyclists frequently switch from the conventional sitting position to a less economical standing posture which allows them to exert more force on the pedals. Pedal frequency, on the other hand, averages ~70 rpm.[4] High mountain ascents are performed by climbing specialists . at intensities close to the AT or ~90% VO2max.[1-3] Because of team requirements, however, some PC are not required to perform maximally during high mountain stages. On the other hand, it should be kept in mind that the top of numerous mountain passes is located at moderate altitude (~2000m). In this regard, it has been reported that highly trained endurance athletes are those who experience more severe gas exchange impairments during acute exposure to hypoxia.[12] Some degree of diffusion limitation could be involved.[12] A recent study has indeed reported radiographic evidence of pulmonary oedema in trained cyclists after high intensity exercise at moderate altitude.[13] 2. Anthropometric Variables Road cycling is a sport that requires performing in a great variety of terrains (i.e. level vs uphill roads) and competitive situations (i.e. individual cycling or drafting behind numerous cyclists). In turn, cycling performance in each of the competition terrains is partly determined by individual morphological characteristics [body mass, height, body surface and frontal areas, body mass index (BMI)].[3] Anthropometric variables might thus greatly differ depending on each PC speciality. TT or flat terrain specialists are usually taller and heavier (180 to 185cm tall, weighing 70 to 75kg, BMI of ~22) than those who excel in uphill climbing (175 to 180cm tall, weighing 60 to 66kg, BMI of 19-20).[3,5] The morphometric characteristics of modern champions being able to excel in both types of terrains are, however, close to those of time trialists (i.e. ~180cm tall, weighing ~70kg).[3] The percentage of body fat, on the other hand, does not significantly differ among the different types of cyclists: starting at values close to 10% (using skinfold techniques)
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during the winter months, it gradually decreases during the season, to reach values around 9% during the spring and close to 8% during 3-week tour races (end of the spring and summer months).[1,14-16] 3. Physiological Variables
3.1 Maximal Power Output and Maximal Oxygen Uptake

Average values of maximal power output attained during an incremental test vary depending on the protocol being used: lower values of 400 to 450W (6.0 to 6.5 W/kg) are recorded during tests with 4-minute increments,[3] whereas power outputs of 450 to 500W (6.5 to 7.5 W/kg) can be obtained during shorter protocols (i.e. 1-minute increments of 25W).[1,5,17-22] Furthermore, maximal power outputs higher than 500W are not unusual in top level time trialists using the latter type of protocol.[5] . Mean values of VO2max reported in PC range between 5.0 to 5.5 L/min or 70 to 80 ml/min/kg when expressed in absolute or relative units, respectively.[1-3,16-25] The highest relative values (~80 ml/min/kg) are found in uphill climbing specialists . (body mass <70kg).[3,5] Thus, high values of VO2max are required for cycling performance at a profes. sional level. Similar values of VO2max are nonetheless found in elite, amateur cyclists of a lower competitive level.[17] Other physiological abilities, such . as the one to maintain high percentages of VO2max (i.e. 90% or more) during prolonged periods of time might have more relevance to success in professional road cycling,[17] as discussed in section 3.2.
3.2 Submaximal Variables
3.2.1 Anaerobic Threshold

Professional cycling racing is a long-duration, high intensity sport and thus requires participants . to possess high AT (i.e. at 90% of VO2max).[1-3,17] There is a great controversy concerning the different methods which can be used to determine the AT in elite athletes (i.e. based on blood lactate determination or ventilatory variables). Some authors have chosen long protocols (4-minute increments) to measure blood lactate levels to determine the exercise
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intensity eliciting a blood level of 4 mmol/L onset of blood lactate accumulation (OBLA)[3] or the individual AT.[2] In contrast, numerous studies from our laboratory have reported the use of ventilatory parameters during shorter, ramp-like protocols (1minute increments) to determine the workload at which the second ventilatory threshold (VT2) or respiratory compensation point (RCP) occurs.[1,5,17,19-22] Keeping in mind the different methodologies used for AT evaluation, most authors agree that the AT of . PC corresponds to ~90% of both VO2max[1,5,17,19-22] and maximal power output.[3,5,22]
3.2.2 Isocapnic Buffering and Hypocapnic Hyperventilation Phases

ison with amateur values could be interpreted as a greater ability to work at high intensities before lactic acid accumulation occurs in the blood. Finally, a greater buffer capacity and a higher tolerance to lactic acidosis is expected from climbing specialists compared with time trialists, given the ability of the former to tolerate repeated bouts of maximum intensity effort during mountain ascents.[5]
3.2.3 Oxygen Uptake Kinetics during Submaximal Exercise

Three physiological gas exchange phases can be identified during rapid incremental exercise testing:[26] phase I, in which CO2 production comes mainly from oxidative metabolism; phase II (isocapnic buffering), during which pulmonary venti. lation (VE) increases in response to the rise in CO2 from buffering with regulation of arterial partial pressure of CO2 (PaCO2); and phase III, in which respiratory compensation for metabolic acidosis with lowering of PaCO2 (hypocapnic hyperventilation or HHV) occurs. The points which limit these 3 phases are denoted VT (between phases I and II), and the aforementioned RCP (between phases II and III). The onset of respiratory compensation of exercise acidosis, when exercise intensity is further increased, marks the final transition from the buffering phase to exercise acidosis. The high workloads at which both VT and RCP . occur in PC (~65% and ~90% of VO2max, respec[1,5,17,19-22] and the marked difference between tively) these values and those recorded in amateur cyclists . (~60% and ~80% of VO2max, respectively),[17,19] suggests that such submaximal variables might be important performance factors in endurance events such as professional road races. On the other hand, no significant differences exist in the isocapnic buffering range of PC and that of elite, amateur cyclists.[19] In contrast, a longer HHV range has been reported in the latter (whether HHV was expressed . in terms of VO2 or W).[19] Both the high values of RCP and the shorter HHV range of PC in compar Adis International Limited. All rights reserved.

. It is well documented that oxygen uptake (VO2) tends to slowly rise during any constant-load cycle ergometer test involving sustained lactic acidosis [i.e. above the lactate threshold (LT) or above VT]: this phenomenon is called the slow component of . VO2 and is typically defined as the continued rise . in VO2 beyond the third minute of exercise.[27] Although several aetiological factors have been proposed (i.e. lactic acidosis, temperature, potassium or cardiorespiratory work), what appears to be the most important factor is a progressive recruitment of less efficient motor units (i.e. type II fibres), with exercise duration.[27] In this regard, we recent. ly studied VO2 kinetics in a group of PC during 20-minute cycle ergometer tests performed at 400W . or ~80% of VO2max.[22] Gas exchange and blood variables and electromyographic (EMG) data from the vastus lateralis muscle were recorded during the tests. The average magnitude of the slow component (7.6 ml/min) was considerably lower than that reported by previous research (22 ml/min) using the same cycle ergometer protocol.[28] Such a finding is suggestive of a great cycling efficiency of PC which is thought to contribute to their renowned ability to sustain high workloads over long periods (i.e. >60 minutes), as shown by previous field studies.[1-3] On the other hand, EMG . data suggested that the primary origin of the VO2 slow component in PC is not attributable to neuromuscular fatigue, at least at intensities up to 80% . of VO2max. These athletes, indeed, exhibit a considerable resistance to fatigue of recruited motor units, at least at submaximal intensities. Such adaptation is probably attained after years of highly demanding training (i.e. ~35 000km per year) as suggested
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in a previous study which compared the physiological response of PC with that of their elite, amateur counterparts.[17] Although further research is needed, the origin . of the VO2 slow component in PC appears to be multifactorial, with a non-negligible contribution of central factors.[22] On the other hand, there seems to exist an inverse correlation between pre-exercise levels of thyroid hormones and the magnitude of the slow component in these athletes.[29] This in turn suggests, at least partly, an involvement of thyroid function on their neuromuscular efficiency during constant load cycling.[29] 4. Cardiovascular Responses and Adaptations
4.1 Heart Rate (HR)
4.1.1 The HR Response to Incremental Exercise: the Conconi Test

(fig. 1), HRd corresponded to a workload (~85% . VO2max) comparable with that eliciting the RCP or the OBLA. The coincidence in time between AT and HR d, does not necessarily imply a causal relationship. Based on echocardiographic evaluations of the participants, those cyclists who showed HRd during incremental exercise were those with greater myocardial wall thickness. It may be speculated that the occurrence of HRd is caused by more efficient cardiac function during high intensity exercise in those athletes who have a greater myocardial wall thickness.[20]
4.1.2 HR as an Indicator of Exercise Intensity

Although a great controversy exists in the literature concerning the so-called Conconi test, previous research by Conconis team and by other authors has confirmed both the existence of a deflection point (HRd) in the heart rate (HR) response to an incremental exercise protocol and its coincidence with the AT in elite athletes (including PC).[30] Indeed, the Conconi test, applicable in field conditions, has become one of the most frequently used exercise tests in sports medicine. Because of the ease of repeated measurements, it is commonly used by elite athletes such as European PC to establish optimum training intensity. We have recently evaluated the applicability of this test for AT determination in 21 PC using a ramplike protocol (workload increases of 25 W/min).[20] The later type of test was chosen following the most recent recommendations made by Conconi et al.[30] for HRd determination. According to Conconis team, the fact that some authors have previously failed to detect HRd may be explained by the protocol used, such as step-like workload increases rather than the more gradual ramp method, which allows increases in HR of <8 beats/min per min of exercise. The HR response showed a deflection point in most PC (~2/3) and was linear in the rest (~1/3). When occurring
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An important issue in professional cycling is to adequately quantify exercise intensity during both training sessions and competitions based on the data of previous laboratory or field incremental tests. In this regard, the blood lactate response during incremental exercise, i.e. the OBLA, is considered to be a good predictor of performance in top-level cycling.[3] Both VT and RCP (which can be determined during progressive exercise testing), have also been shown to be important determinants of performance and fitness level in endurance exercise, especially the RCP.[1] This, in turn, permits the quantification of training and competition in different phases in terms of exercise intensity, i.e. phase . I or low intensity (<VT or <65% of VO2max), phase II or moderate intensity (between VT-RCP or be. tween 65 to 90% of VO2max) and phase III or high . intensity (>RCP or >90% of VO2max).[31] On the other hand, recent technological developments have made it possible to measure power output (W) on a bicycle with a power measuring device (i.e. the Schoberer Rad Messtechnik Training System) and thus to prescribe training loads based on power output eliciting the aforementioned ventilatory or lactate markers. Moreover, power output may be the most direct indicator of exercise intensity in general.[32] During actual cycling, however, power output is much more variable than HR,[32] which limits its use for training prescription in PC. To date, HR is the most frequently used parameter for evaluating the level of intensity attained during training sessions and competition in profesSports Med 2001; 31 (5)

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190 180 170 160

HR (beats/min)

HRd = 164 beats/min

y = 0.163x + 97.952 (r = 0.984)

150 140 130 120 110 100 90 200 300 Workload (W) 400 500

y = 0.274x + 53.147 (r = 0.972)

Fig. 1. Determination of the point of heart rate deflection (HRd) in 1 professional cyclist (former world champion) from the heart rate

(HR)/workload (W) relationship. At times (such as in this example), the initial data points (i.e. below 100 to 110 beats/min) do not fall on the straight line and should be ignored according to the methodology of Conconi and co-workers.[30] (From Lucia et al.,[20] with permission).

sional cycling.[21] When training is based on HR data, training orientation does not necessarily require periodic readjustment of target power output (i.e. at LT or RCP) by repeated testing during the season. Indeed, it has been recently shown that target HR values corresponding to performance markers such as LT or RCP remained stable (~155 and ~175 beats/min, respectively) during the course of a complete sports season in a group of 13 PC, despite a significant improvement in performance throughout the training season (i.e. increases in the power output eliciting LT or RCP).[21] Comparable findings were also obtained by Hoogeven,[14] that is, no change throughout the season in HR values corresponding to VT. Interestingly, maximal HR does not significantly change during the season, whereas recovery values (recorded at 3 and 5 minutes after exercise) show a consistent decrease.[21] Although speculative, the decrease in post-exercise HR might be attributed to an increase in stroke volume or to
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a decrease in sympathetic tone, both of which are well known adaptations to endurance training.[33]
4.2 Cardiovascular Adaptations
4.2.1 Cardiac Adaptations

Some studies have reported echocardiographic parameters in PC.[20,34-40] Cardiac hypertrophy is not an uncommon finding among these athletes. For example, Rodrguez Reguero et al.[38] found the existence of ventricular hypertrophy (i.e. diastolic ventricular thickness 13mm and a mean ventricular mass index of 152 g/m2) in 21 participants from a group of 40 PC (mean age of 26 years and 3 to 10 years of competition experience in the professional category). Further research from the same group suggested that, when existing, cardiac hypertrophy is a physiological adaptation to strenuous exercise (i.e. 30 000km cycled per year) rather than a pathological condition related to haemodynamic or endocrine disturbances.[39] A study from our laboratory has also reported echocardiographicallySports Med 2001; 31 (5)

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derived cardiac dimensions in 21 top-level PC, including world champions and winners of major 3week tour races (mean age of 25 years and mean competition experience in the category of 4 2 years).[20] We found a lower degree of ventricular hypertrophy (i.e. mean ventricular mass index of ~116 g/m2) compared with the study by Rodrguez Reguero et al.[38] Our data were similar to those reported in previous studies for highly trained endurance athletes in general,[41-44] that is, PC show a cardiac profile expected of endurance athletes with predominantly eccentric left ventricular hypertrophy (enlarged left ventricular end-diastolic internal diameter and proportional increase in wall thickness). The incidence of electrocardiographic abnormalities (i.e. ST-T segment alterations) is very low among PC.[34,38] Even extreme endurance events such as 3-week races do not appear to have a deleterious effect on the hearts of PC. In a study conducted during the Giro dItalia, Bonetti and co-workers showed no evidence of myocardial damage throughout the 3-week period in any of the participants using a marker of myocardial ischaemia such as cardiac troponin T.[45]
4.2.2 Vascular Adaptations

aerodynamic cycling positions.[51] Surgical mobilisations or reconstructions of the iliac arteries are possible types of treatment.[51] Although long term follow-up reports are lacking, some Spanish PC have performed successfully after surgery (unpublished data). 5. Pulmonary Responses and Adaptations Pulmonary volumes of PC do not differ from those of amateur, elite counterparts (i.e. vital capacity of ~6.0L for a height of 175 to 180cm).[24] . Similarly, maximal values of VE do not differ between PC and amateurs (180 to 190 L/min in both groups).[17,24,52] In contrast, the breathing pattern of the former during incremental exercise shows 2 unique characteristics, compared with other well trained endurance athletes (including amateur cyclists):[53-55] (i) a prolongation of expiratory duration, especially at high workloads; and (ii) a lack of tachypnoeic shift at high exercise intensities, that is, tidal volume (VT) does not show a plateau at near-maximal intensities.[24] Both characteristics might be interpreted as an adaptation to the demands of professional cycling.[24] The fact that PC do not adopt the tachypnoeic shift can be viewed as a particularly efficient respiratory adaptation to training from both metabolic and mechanical points of views.[24] 6. Physiological Adaptations during a Sports Season The prevailing specific adaptations to training incurred by already well-trained endurance athletes is, in general, a controversial issue. Four prospective reports have evaluated the physiological adaptations which occur in PC as a response to training and competition during the main part (~7 months) of a typical sports season.[14,19,56,57] The sports season of PC generally includes 3 different periods in terms of training volume and/or intensity, i.e. precompetition or training (winter), competition (spring and summer, usually with an active rest period in between) and postcompetition or active rest periods (fall) [table II].[14,19,56,57] During 2 to 3 weeks
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Professional cycling also has an effect on large arteries. Abergel et al.[40] found a 13% larger diameter and mean diastolic intima-media thickness in the carotid arteries of 149 participants in the Tour de France compared with poorly trained controls. The work from the research group of Chevalier[46-49] in Lyon, France, provides some insight into the prevalence of flow limitation in the iliac arteries of PC. Of the 223 operations that Chevalier has reported from 1985 to 1996 in the ilio-femoral region, 63% were performed on well trained cyclists (15% were PC).[50] An average of 120 000km must usually be cycled before the rider develops complaints, consisting mainly of leg pain and a powerless feeling in the legs.[51] Flow limitation is attributable to either intravascular (endofibrosis) or functional lesions, i.e. kinking and/or excessive lengthening of the iliac arteries over the ventral surface psoas muscle during hip flexions, especially when the rider adopts
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of the last period, PC adopt an almost sedentary lifestyle.

6.1 Maximal Variables

Table II. Training characteristics of professional cyclists during the year[19,21,56] Rest (fall) Average weekly training (km) Exercise intensitya low (%) moderate (%) high (%) a ~88 ~11 ~1 ~78 ~17 ~77 ~15 ~270 Precompetition Competition (winter) (spring-summer) ~700 ~800

. Some authors have shown that the VO2max of elite cyclists increases during the season.[14,58,59] The data from our laboratory,[56] however, are in line with those of most previous studies which showed no significant effects of training intervention on . VO2max in well-trained athletes such as runners,[60] swimmers,[61] and elite amateur cyclists.[62] Further, we have detected no difference in maximal power output throughout the season.[19,56] Based on these findings, it seems that once a certain training status is reached (i.e. professional cycling category), further increases in training intensity and volume are no longer associated with improvements . in VO2max. It would appear that other physiological characteristics, such as the ability to maintain high . percentages (i.e. 90%) of VO2max during prolonged periods (>30 minutes), play a more relevant role in successful endurance cycling. It is consequently felt that training programmes should be designed to improve this ability.
6.2 Submaximal Variables: Metabolic and Neuromuscular Adaptations

. VO2max = maximal oxygen uptake.

~5 ~8 . Exercise intensity: low (<65% VO2max), moderate (65-90% . . VO2max), high (>90% VO2max).

given workload),[56] nor in the isocapnic buffering range.[19] In contrast, Weston et al.[62] reported that the racing performance of well-trained amateur cyclists might increase during the season mainly as the result of improved muscle buffering capacity. Such a finding, however, might not be directly extrapolated to PC with a higher fitness level and a more solid training background. 7. Endocrine System It has been documented that some endurancetrained men, especially runners, might exhibit some subclinical alterations in their hypothalamic-pituitarytesticular (HPT) axis which are partly attributable to a hyperactivation of the hypothalamic-pituitaryadrenal (HPA) axis.[63] Such alterations might include decreases in basal levels of testosterone,[64] or in sperm quality.[65] Although subclinical decreases in basal levels of testosterone have also been reported in PC after an intense training period reflecting a certain catabolic state (i.e. decreased testosterone : cortisol ratio), such changes do not seem to affect cycling performance.[15] Moreover, a previous longitudinal study did not show any significant change in the HPT (including sperm characteristics) and HPA axes of 12 PC over a full sports season.[16] Although sperm motility was transiently decreased during the competition period in most participants, such an alteration was attributed solely to physical factors, i.e. testicular and/or prostatic microtrauma against the saddle or increased intrascrotal temperature. Scarce data are available in the
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In a recent study,[57] we found no overall training effect throughout the season in the ventilatory . response of 13 PC during a ramp-protocol (i.e. VE, VT, ventilatory equivalents for O2 and CO2, timing of respiration, etc.), despite a significant improvement in performance (i.e. increases in both the . power output and the percentage of VO2max eliciting RCP). In contrast, several metabolic and neuromuscular variables showed major changes during the season in the same participants, such as: (i) lower circulating lactate levels and possibly increased reliance on oxidative metabolism at a given submaximal intensity (up to 400W); and (ii) an enhanced recruitment of motor units in active muscles, as suggested by EMG data.[56] No significant improvements, however, were found in the cyclists buffering capacity during the season (i.e. no significant changes in blood pH and HCO3- levels for a
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literature concerning the adaptations of other hormonal systems to professional cycling. It has nevertheless been reported that the resting activity of the renin-angiotensin-aldosterone system is normal in these athletes.[39] 8. Haematological Variables and Blood Doping Sports haematology and blood doping have become a contentious topic in elite sports in the last 3 decades. Since recombinant human erythropoietin (r-HuEPO) is demonstrably effective in increas. ing haemoglobin levels, VO2max and physical work capacity,[66-68] the lack of a reliable test (until most recently) to confirm its use may have presumably induced many PC to experiment with this drug over the last 15 years. In fact, the possible health risks of hyperviscosity and thrombogenicity associated with the misuse of this drug could have caused the mysterious deaths of some European riders between 1987 and 1990.[69] The suspected association between elite cycling and blood doping with r-HuEPO was confirmed by both the discovery of vials of this drug in a car belonging to a professional cycling team during the 1998 Tour de France and the recent finding of abnormally high erythropoietin levels in several frozen urine samples collected during the aforementioned race.[70] To dissuade the use of r-HuEPO and to minimise the health risks associated with the abuse of this drug, the International Cycling Union has imposed an upper limit of 50% on haematocrit levels since 1997. Disadvantages of using this threshold however, include: postural effects,[71] ease of manipulations through interventions such as saline infusion,[68] and large natural variations among individuals.[68,72] Concerning the latter, it has been recently documented that a subset of endurance athletes including elite cyclists (i.e. 2 to 8% of total) might naturally surpass the 50% threshold.[73-75] In fact, some authors have suggested that those athletes with high haematocrit levels (i.e. close to 50%) might be more genetically predisposed to perform successfully in endurance sports.[73] Other authors, in contrast, have reported haematocrit levels consistently below 50%
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(mean of 43.0 0.02%; range 0.39 to 0.48) in 353 blood samples collected from PC during 1980 to 1986, before r-HuEPO was commercially available.[76] The later finding is in agreement with the prevailing idea in sports haematology, that is, dilutional pseudoanaemia secondary to plasma volume expansion is a common finding among endurance athletes.[77] Fortunately, both indirect (based on markers of altered erythropoiesis)[68] and direct methods[70] recently reported in the biomedical literature are already available for identifying current or recent users of r-HuEPO. In this regard, it must be emphasised that in a number of laboratory studies from which several conclusions were drawn about the physiology of professional road cycling . (i.e. analysis of the VO2 response), haematocrit levels were reported to be below 50% in all the participants.[5,22,29] The fact that blood doping appears to work in PC indirectly supports the idea that top performance in this sport is limited by central (cardiopulmonary system) rather than by peripheral factors (oxygen utilisation by working muscles). Indeed, the only potential detrimental effect of this method in terms of performance is decreased stroke volume secondary to increased blood viscosity,[77] but actually its effect appears to be negligible, at least at haematocrit levels below 50 to 55%. 9. Nutritional Habits In general, PC might be defined as a homogeneous group with a similar nutrition intake, eating habits and nutritional needs.[78,79]
9.1 Tour Races

Few studies have evaluated the feeding pattern of PC during 3-week tour races, in both actual[23,79] and simulated conditions.[80,81] During these races, average daily energy intake is as high as 23 to 25MJ.[23,79] One of the main concerns in these types of events is to replenish bodily glycogen stores within 18 hours (from the end of each daily stage, at ~5pm, to the beginning of the next one, at ~12pm) over a 3-week period. Although reports with muscle biopsies are not available during acSports Med 2001; 31 (5)

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tual cycling races, cyclists carbohydrate intake during these extreme events appears to be sufficient (~60% of total caloric intake or >800g/24h period or 12 to 13 g/kg bodyweight per day),[23,79] based on the recommendations of previous research.[82,83] In particular, the high carbohydrate intake during the first 6 hours after the race (1.1 g/kg bodyweight) is crucial.[79] Additionally, an interesting factor for muscle glycogen resynthesis is the addition of protein (0.35 g/kg bodyweight) to carbohydrate in the first hour after exercise.[79] The latter is achieved by consuming prepared semi-solid foods consisting basically of a mixture of cereals, dairy products and fruits. Carbohydrate intake during exercise, however, is rather low (average 25 g/h),[79] and below the recommendations (30 to 60 g/h) for maintenance of a high rate of carbohydrate oxidation during prolonged strenuous exercise.[83] Another remarkable characteristic is the high protein intake (approximately 3 g/kg bodyweight per 24 hour period),[23,79] clearly above that recommended for endurance athletes in general (1.2 to 1.4 g/kg bodyweight per day).[84] This protein intake is mainly because of the high overall energy intake, since the relative contribution of protein to energy is not higher than 14 to 18%.[23,79] On the other hand, the diet of PC during tour races provides 23 to 25% of energy from fat, with biscuits and confectionery providing as much as 27 to 43% of this energy substrate.[23,79] There exists considerable variation regarding the fluid intake reported during tour races, i.e. 3.3L per 24 hours in the study by Garca-Rovs et al.[79] versus 6.7L per 24 hours in that by Saris et al.[23] Such variability comes mainly from the carbohydrate feeding pattern adopted during exercise, since the participants in the study by Saris et al. consumed a large amount of carbohydrates in the form of sports drinks (4L per stage).
9.2 Training

tein intake during the latter.[78] Vitamin intake during both training and competition periods seems to be well above the recommended daily amount for healthy adult males, especially when considering that most riders consume vitamin supplements.[23,78] 10. Conclusion To date, some descriptive studies have analysed the physiological responses and adaptations to such an extreme endurance sport as professional road cycling.[1-6,9,10,14-22,24,29,34,36-39,45,56,57] Furthermore, a great amount of data have been collected during actual races. As a result, the body of knowledge concerning cycling physiology has considerably increased in the last decade. For instance, we know that the results from previous research with elite, amateur cyclists, might not be directly extrapolated to professional riders. Further nondescriptive research is however needed in the field. Given the great performance level of the participants, this sport could serve as a model to better understand the mechanisms involved in the human responses/adaptations to endurance exercise. Acknowledgements
We are indebted to Asociacin Deportiva Banesto for supporting our research.

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Correspondence and offprints: Dr Alejandro Luca, Departamento de Ciencias Morfolgicas y Fisiologa (Edificio A, Despacho 330), Universidad Europea de Madrid, 28670 Madrid, Spain. E-mail: alejandro.lucia@mrfs.cisa.uem.es

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