The Relationship Between Freely Chosen Cadence
Transcription
The Relationship Between Freely Chosen Cadence
International Journal of Sports Physiology and Performance, 2012, 7, 375-381 © 2012 Human Kinetics, Inc. www.IJSPP-Journal.com ORIGINAL INVESTIGATION The Relationship Between Freely Chosen Cadence and Optimal Cadence in Cycling Umberto Emanuele, Tamara Horn, and Jachen Denoth Purpose: The main aim of this study was to compare the freely chosen cadence (FCC) and the cadence at which the blood lactate concentration at constant power output is minimized (optimal cadence [Copt]). The second aim was to examine the effect of a concomitant change of road incline and body position on FCC, the maximal external power output (Pmax), and the corresponding Copt. Methods: FCC, Copt, and Pmax were analyzed under 2 conditions: cycling on level ground in a dropped position (LGDP) and cycling uphill in an upright position (UHUP). Seven experienced cyclists participated in this study. They cycled on a treadmill to test the 2 main hypotheses: Experienced cyclists would choose an adequate cadence close to Copt independent of the cycling condition, and FCC and Copt would be lower and Pmax higher for UHUP than with LGDP. Results: Most but not all experienced cyclists chose an adequate cadence close to Copt. Independent of the cycling condition, FCC and Copt were not statistically different. FCC (82.1 ± 11.1 and 89.3 ± 10.6 rpm, respectively) and Copt (81.5 ± 9.8 and 87.7 ± 10.9 rpm, respectively) were significantly lower and Pmax was significantly higher (2.0 ± 2.1%) for UHUP than for LGDP. Conclusion: Most experienced cyclists choose a cadence near Copt to minimize peripheral fatigue at a given power output independent of the cycling condition. Furthermore, it is advantageous to use a lower cadence and a more upright body position during uphill cycling. Keywords: FCC, uphill, level ground, body position, power output On modern multigeared bicycles, the cyclist is free to choose any available gear ratio to achieve a target cycling velocity. The choice of the gear ratio at a given cycling velocity in turn determines the cadence. Thus, as noted by Vercruyssen and Brisswalter,1 the cadence is one of the only variables that an athlete can modify during cycling exercise to optimize performance. Hence, cadence selection has been of interest since the multigeared chain-driven bicycle was invented in the late 19th century. For more than 100 years, cyclists, coaches, and researchers have debated over cadence choice. Even in the last 10 years, several scientific studies on the freely chosen cadence (FCC) have been published, further demonstrating that there is a continuous discussion on the theory and practice of cycling. Research has shown that FCC is a highly individual characteristic and is affected by several “external” factors.2 It has been reported that external power output (Pext),3 road gradient,4 crank inertial load,3 drafting,5 fatigue,6 training,7 muscle-fiber-type composition,8 and circadian rhythm9 influence FCC. Despite this intense discussion, the underlying factors leading to the choice of a particular cadence are still a subject of debate. Theoretically, cyclists should choose an “optimal cadence” at which relevant physiological The authors are with the Institute for Biomechanics, ETH Zurich, Zurich, Switzerland. and/or biomechanical variables are optimized. However, several studies document the multitude of definitions for “optimal cadence” in cycling (for a review see, eg, Marais and Pelayo10). The term optimal cadence has been defined as the cadence with the lowest EMG activity for a given Pext,11 the cadence that minimizes the summed joint moments,12 the cadence that produces the lowest neuromuscular fatigue,13 the cadence with the highest Pext at a specific performance level,14 and the most efficient cadence for energy expenditure.15 The “cadence paradox,”16 which states that competitive cyclists and trained subjects choose a higher cadence than the most efficient cadence (Ceff), has been an especially interesting topic in the past.17 Of most interest (at least for a competitive road cyclist) is the cadence that produces the best performance to win a race, not Ceff. Therefore, the optimal cadence (Copt) is defined here as the cadence that corresponds to the apex of the Pext–cadence relationship (Pmax) at a performance level that can be sustained for the given task.14 From a theoretical perspective, cyclists should choose a cadence near Copt to maximize Pext for the given task or to minimize peripheral fatigue at a given Pext. Results from previous studies showed that both FCC3 and Copt18 were significantly lower during uphill cycling than during cycling on level ground with the same body position. Taken together, these results suggest that both FCC and Copt are affected in a similar way by road incline, but no 375 376 Emanuele, Horn, and Denoth study has addressed whether the difference between FCC and Copt is the same independent of road incline. Moreover, to the best of our knowledge, no study has compared FCC and Copt at a submaximal performance level. A study by Emanuele and Denoth18 showed that the road incline not only decreased Copt but also decreased Pmax. In addition, that study showed that the change in body position from a dropped posture (DP) with the cyclist’s hands on the drops of the handlebars and arms fully extended to an upright posture (UP) with the cyclist’s hands on the top portion of the handlebars and arms fully extended significantly increased Pmax without affecting the corresponding Copt. These results led to the conclusion that Copt and FCC should be lower and Pmax should be higher for uphill cycling with UP than for level-ground cycling with DP.18 However, the influence of a simultaneous change in road incline and body position as seen in field observations of cyclists has not been analyzed in an experimental study. The main aim of this study was to compare FCC and Copt at an endurance-performance level that can be sustained for approximately 1 hour. The hypothesis was that at a performance level corresponding to the maximal lactate steady state (MLSS), experienced cyclists would choose an adequate cadence close to Copt independent of the cycling condition. The second aim of this study was to examine the effect of a concomitant change in road incline and body position on FCC, Copt, and Pmax at an endurance-performance level corresponding to MLSS. The hypothesis was that FCC and Copt would be lower and Pmax higher during uphill cycling with UP than during level-ground cycling with DP. Methods Subjects Seven well-trained male amateur cyclists (31 ± 6 y, 182.0 ± 4.5 cm, 71.7 ± 6.7 kg) who have competed at the national level volunteered to participate in this study. They were all informed of the nature of this study, as well as the potential risk and discomfort associated with the experimental procedures, before they gave their written consent to participate. The ethical committee of ETH Zurich approved the experimental design. Determination of FCC. To determine FCC during level-ground cycling (FCCLG) the subjects cycled with DP at a road inclination of 0% and a speed of 30.2 km/h. During this test, the cyclists were encouraged to change gears and use the gear that felt the most comfortable. However, because of the limited gear ratios, there were limitations on the FCCs. Only the following cadences could be chosen: 60, 65, 70, 75, 80, 85, 90, 95, 100, and 105 rpm. After a 10-minute warm-up at 100 W, Pext was increased until it reached a level that the cyclist thought he could maintain for approximately 1 hour. This subjective estimation of PMLSS was reached within 5 to 7 minutes. Pext and gear were then held constant for 1 minute, and the cadence (FCC) and Pext were recorded. The subjects were then given a 20-minute rest period. To determine FCC during uphill cycling (FCCUH), the same protocol was used, except that subjects cycled with UP on the treadmill with an inclination of 7% and a speed of 15.1 km/h. Lactate Minimum Test. The determination of the lactate minimum power output (PLM) using a lactate minimum test (LMT) is a reliable and valid method to predict PMLSS and therefore a good predictor of the specific endurance performance level that can be sustained for approximately 1 hour.19,20 During the LMT, the participants cycled with UP and a constant cadence of 80 rpm, which corresponds to the mean Copt measured by Emanuele and Denoth18 during cycling using an ergometer. The method used to determine PLM was based on Fontana et al21 (Figure 1). Briefly, the LMT consisted of 2 incremental tests. The first incremental test started at 100 W and increased 50 W every minute until exhaustion to induce lactic acidosis, followed by 1 minute at 100 W and 7 minutes of complete rest (part 1). After the rest period, a second incremental test started at 100 W with an increase of 25 W every 90 seconds (part 2). During the LMT, heart rate, oxygen uptake, carbon dioxide output, minute ventilation, and breathing frequency were continuously recorded. Blood lactate concentration (bLa) was measured at rest, at exhaustion in part 1, during recovery in part 1 (after 2, 4, Experimental Design The subjects were asked to attend 3 test sessions within a 3-week period with at least 2 days between single test days. To improve the reliability of the measurements, the participants were asked to control a number of variables; for the details see Emanuele and Denoth.14,18 The purpose of the first test session was to determine individual FCC and estimate the individual Pext at maximal lactate steady state (PMLSS). The purpose of the second and third test sessions was to estimate Copt and Pmax at MLSS when cycling on level ground with DP and when cycling uphill with UP, respectively. Figure 1 — Schematic of the protocol for the lactate minimum test. The test consists of 2 incremental tests, separated by 1 minute at 100 W and 7 minutes of complete rest. The first incremental test started at 100 W with 50-W increments every minute until exhaustion to induce lactic acidosis. The second incremental test started at 100 W with 25-W increments every 90 seconds. × = Measurements of blood lactate concentration (bLa). FCC and Optimal Cadence in Cycling 377 and 7.5 min), and at the end of each stage in part 2. The bLa levels during part 2 of the LMT were plotted against power output, and a third-order polynomial curve was fitted through these data points.21 The PLM corresponds to the “lactate minimum” in this polynomial curve. PLM was used as an estimation of PMLSS at 80 rpm, the exercise intensity needed for the second and third test sessions. Determination of Copt. To determine Copt at MLSS during uphill cycling, the subjects performed 3 incremental exercise tests at the following 3 cadences: FCCUH – 10 rpm, FCCUH, and FCCUH + 10 rpm in a randomized order. During these tests, the subjects cycled with UP on the treadmill at an 7% inclination at 15.1 km/h. At each cadence, a 2-minute warm up at 100 W was followed by an incremental exercise test comprising 4 increments in Pext (Figure 2): P80MLSS – 20 W (3 min), P80MLSS – 10 W (3 min), P80MLSS (2 min), and P80MLSS + 10 W (2 min). The incremental test was followed by a 2.5minute cooldown at 100 W. Between the 3 incremental exercise tests, the subjects were given a 14-minute rest to prevent fatigue. bLa was measured at the beginning of the warm-ups and at the end of each stage of the incremental exercise tests. To determine Copt at MLSS, the increase in bLa was analyzed. The increase in bLa from rest until the end of the exercise test was plotted against the cadence used (Figure 3[a]). A second-order polynomial-regression line was fitted through these data points to determine the bLa–cadence relationship. Copt at MLSS corresponds to the “lactate minimum” in this polynomial curve. A quadratic curve constrained to pass through the origin was then fit to Copt at MLSS and PMLSS at 80 rpm to assess the individual Pmax at MLSS (Figure 3[b]).14 To determine Copt at MLSS during level-ground cycling, the same protocol was used, except that the treadmill was set at an inclination of 0%, the speed was increased to 30.2 km/h, and the participants cycled with DP. The bLa–cadence relationship for level-ground cycling was then compared with the bLa–cadence relationship for uphill cycling. Pext for uphill cycling with UP corresponding to the intersection point of the 2 bLa– cadence relationships was used to fit a quadratic curve constrained to pass through the origin to assess the individual Pmax at MLSS for level-ground cycling with DP. Figure 3 — (a) Blood lactate concentration (bLa) in relation to cadence. Mean values for bLa at freely chosen cadence (FCC), FCC – 10 rpm, and FCC + 10 rpm are shown for uphill cycling (white triangles) and for level-ground cycling (white squares). Second-order polynomial-regression curves were fitted to the data on level ground cycling with the dropped posture (LGDP; dashed line) and uphill cycling with the upright posture (UHUP; solid line) to assess the optimal cadence (Copt) for UHUP (black triangle) and Copt for LGDP (black square). The cadences were normalized relative to the Copt at maximal lactate steady state (MLSS) for LGDP. (b) Power–cadence relationships at the performance level corresponding to the MLSS. To assess the maximal power output (Pmax) at MLSS for UHUP, a quadratic curve (solid line) constrained to pass through the origin was fitted to the assessed Copt for UHUP (black triangle) and the power output at MLSS estimated with the lactate minimum test at 80 rpm (P80MLSS, white triangle). To assess Pmax at MLSS for LGDP (black square) the Pext for UHUP corresponding to the intersection point of the 2 bLa–cadence relationships (white square) was used to fit a quadratic curve constrained to pass through the origin (dashed line). These power outputs and cadences were normalized relative to Pmax at MLSS and the corresponding optimal cadence (Copt at MLSS) for LGDP. Equipment Figure 2 — Schematic of the protocol for test sessions 2 and 3. PMLSS80 = power output at maximal lactate steady state estimated with a lactate minimum test at 80 rpm. × = Measurements of blood lactate concentration (bLa). In all tests, the subjects exercised on a normal road-racing bicycle with 24-36-48 teeth chain rings and 12-13-1415-17-18-19-20-21-24 teeth cog set. The bicycle was adjusted so that the vertical and horizontal positions of the saddle and the handlebar related to the crank axis matched each subject’s own bicycle. The racing bicycle was equipped with a standard crank (length = 170 mm), clipless pedals, and a professional (8 strain gages) SRM PowerMeter (Schoberer Rad Messtechnik, Jülich, Germany). Using the SRM PowerMeter, Pext, cadence, and heart rate were recorded. During all tests except the LMT, the bicycle was mounted on a treadmill (Woodway, Weil 378 Emanuele, Horn, and Denoth am Rhein, Germany). Thereby, the bicycle was fixed with the fork to a sliding carriage, which allowed a horizontal bicycle translation relative to the laboratory (Figure 4). Pext was adjusted by changing the mass of a weight magazine. This magazine was connected to a wire that ran over a pulley placed behind the treadmill and was then tied to the back of the bicycle.3,22 The mass needed to achieve the target Pext was calculated for each subject based on his measured Pext without the weight magazine. During the LMT, the bicycle was mounted on an indoor trainer (Flow, Tacx, Wassenaar, Netherlands). The gas-exchange and ventilatory variables were measured breath by breath with an Oxycon Mobile device (Viasys Healthcare, Höchberg, Germany). The lactate concentration in the blood samples (20 μL) taken from the earlobes was analyzed with a Biosen C-Line analyzer (EKF Industrie-Elektronik, Barleben, Germany). Statistics All of the statistical analyses were performed using SPSS Statistics 17 (SPSS Inc, Chicago, IL). The level of significance was set at P < .05. Based on the estimated coefficient of variation for the measured bLa of 5%, the 95% confidence interval (CI) for assessing the individual Copt at MLSS (Figure 5) was calculated using the modelbased residual bootstrapping method for regression.14,18 Assuming that the cyclists use the next higher (Copt+) or lower (Copt–) possible cadence to the real Copt (Copt*) (Figure 5), the theoretical 95% CI for the difference between FCC and the measured/estimated Copt at MLSS (ΔC) was calculated using the 95% CI for the individual Copt at MLSS and the available gear ratios. The theoretical 95% CIs for ΔC were also calculated assuming that the cyclists chose Copt+ + 5 rpm or Copt– – 5 rpm. The statistical power for detecting a relevant ΔC over 5 rpm was calculated for a sample size of 7. All the parameter values were compared using the Student paired t test. The variables were summarized with descriptive statistics (mean ± SD). Figure 4 — The standard bicycle mounted on the treadmill. The fork was fixed to the sliding carriage, which allowed horizontal bicycle translation relative to the laboratory (30 cm). Figure 5 — Schematic of the different cadences. Black squares mark the available cadences that the cyclist can choose. The white circle marks the cyclist’s freely chosen cadence (FCC). The asterisk (*) marks the real optimal cadence (Copt*). The black triangle marks the measured/estimated optimal cadence (Copt). Copt– is the next lower available cadence to Copt*. Copt+ is the next higher available cadence to Copt*. ΔC is the difference between FCC and Copt. The possible distribution of Copt using the method to determine Copt described in this study is shown (thin solid line). The 95% confidence interval (CI) to assess Copt is marked by the dotted lines. Results Based on the coefficient of variation for the measured increase in bLa, the residual bootstrap yielded a 95% CI for assessment of the individual Copt at MLSS of 4.8 rpm (Figure 6[a]). Assuming that the cyclists use Copt+ or Copt–, the theoretical 95% CI for ΔC was 0 ± 7.4 rpm (Figure 6[b]). During level-ground cycling, ΔC of all cyclists was within this CI, and during uphill cycling the difference of 1 cyclist was outside this CI. Assuming that the cyclists use Copt+ + 5 rpm or Copt– – 5 rpm, the theoretical 95% CIs for ΔC were 7.5 ± 3.7 rpm and –7.5 ± 3.7 rpm. During level-ground cycling, ΔC of 6 cyclists was outside this CI, and during uphill cycling, ΔC of 4 cyclists was outside this CI. Thus, during level-ground cycling, 6 cyclists used Copt+ or Copt– and 1 cyclist used Copt+ or Copt+ +5 rpm. During uphill cycling, 4 cyclists used Copt+ or Copt–, 1 cyclist used Copt– or Copt– – 5 rpm, and 1 cyclist used Copt+ + 5 rpm. For uphill cycling with UP and level-ground cycling with DP, the determined FCC and Copt at MLSS were not significantly different. The statistical power to detect a relevant difference of 5 rpm between FCC and Copt at MLSS was .84 for a sample size of 7. FCC at the subjective self-selected PMLSS (278 ± 30 W) was significantly lower (P < .05) when cycling uphill with UP (82.1 ± 11.1 rpm) than when cycling on level ground with DP (89.3 ± 10.6 rpm). PMLSS at 80 rpm estimated using the LMT was 261 ± 24 W. The assessed individual Copt at MLSS was significantly higher (P < .01) for level-ground cycling with DP (87.7 ± 10.9 rpm) than for uphill cycling with UP (81.5 ± 9.8 rpm). Copt at MLSS for level-ground cycling was 7.5% ± 2.3% higher than for uphill cycling. Finally, Pmax at MLSS was significantly higher (2.0% ± 2.1%; P < .05) for uphill cycling with UP than for level-ground cycling with DP (Figure 3[b]). FCC and Optimal Cadence in Cycling 379 Figure 6 — (a) The relationship between optimal cadence (Copt) and freely chosen cadence (FCC). The data for uphill cycling with the upright posture (white triangles) and level-ground cycling with the dropped posture (white squares) are shown for all subjects (n = 7). The solid line represents the identity line (FCC = Copt). The 95% confidence interval (CI) for assessing Copt is shown by the 2 dashed lines. The dotted lines mark the available cadences that the cyclist could choose. (b) The individual differences between FCC and Copt (ΔC). The ΔCs for uphill cycling with the upright posture and level-ground cycling with the dropped posture are shown for all subjects (n = 7). The solid lines mark the theoretical 95% CI for ΔC assuming that the cyclists choose the next higher (Copt+) or lower (Copt–) possible cadence to the real Copt. The dashed lines mark the theoretical 95% CIs for ΔC assuming that the cyclists choose the next higher Copt+ + 5 rpm or Copt– – 5 rpm. The gray zone marks the intersection of 2 CIs. For the data in this gray zone it is not possible to determine if the cyclists used Copt+ or Copt+ + 5 rpm or Copt– or Copt– – 5 rpm. Discussion FCC and Copt The main purpose of this study was to compare FCC and Copt at an endurance-performance level that corresponded to MLSS. The hypothesis was that experienced cyclists would choose an adequate cadence close to Copt independent of the cycling condition. The results revealed that most but not all experienced cyclists choose the next higher (Copt+) or the next lower (Copt–) possible cadence to Copt, independent of the cycling condition. However, the results also showed that, independent of road incline and body position, FCC and Copt at MLSS were not significantly different. Here it must be stated that the statistical power to detect a difference between FCC and Copt at MLSS was greater than .8 only for an assumed difference of 5 rpm between FCC and Copt at MLSS. On the other hand, a difference between FCC and Copt at MLSS of less than 5 rpm is not relevant from a performance-related point of view, because such a difference reduces Pext less than 0.5%.14 Furthermore, most of the individual differences between FCC and Copt at MLSS can be attributed to the discrete available gear ratios and the precision in determination of Copt at MLSS. According to Emanuele and Denoth 14 C opt is defined as the cadence that corresponds to the apex of the Pext–cadence relationship at a specific performance level, for example, at a fixed bLa. From a theoretical perspective, Copt can be indirectly assessed by measuring different variables at a constant Pext while manipulating the cadence. One of these indicative variables is bLa. Emanuele and Denoth14 showed that to assess Copt at a fixed bLa, the Pext–cadence relationship can be fitted using a quadratic-regression line that is constrained to pass through the origin. This result implies that Copt can also be assessed using a quadratic bLa–cadence relationship at a constant Pext. Thus, the results from this study confirm the assumption that most competitive cyclists choose a cadence near Copt to minimize peripheral fatigue at a given Pext or to maximize Pext for the given task. This assumption has been stated by different authors but has never been experimentally demonstrated. The following additional indicative variables for Copt at a constant Pext have been analyzed previously: neuromuscular fatigue,13 EMG activity,11,23 and time to exhaustion.24 All of the single data sets of these studies were well fitted with a second-order polynomial-regression curve (R2 = .88–.99) to assess Copt. However, none of these studies assessed Copt and compared it with FCC. Thus, this is the first study to show that most competitive cyclists choose an adequate cadence close to the individual Copt at a specific endurance-performance level. Furthermore, the comparison between level-ground cycling with DP and uphill cycling with UP revealed that not only interindividual differences in FCC but also intraindividual differences in FCC are related to Copt. These are experimental indications of a causative relationship between FCC and Copt. This relationship can also be deduced from the observation that both FCC6 and Copt25 decrease 380 Emanuele, Horn, and Denoth with increasing fatigue. Thus, the cadence paradox16 is resolved because most competitive cyclists choose a cadence near Copt, which is clearly a higher cadence than the most efficient cadence. Kohler and Boutellier16 have already explained that FCC exceeds Ceff. In their theoretical study, they concluded that FCC is not fixed but depends on the duration of the race and ranges between Copt and Ceff. However, in their conclusion they neglected the fact that not only FCC but also Copt and Ceff depend on performance level or, rather, race duration.14,26 Even their own calculation showed that Copt, Ceff, and the difference between these cadences depend on performance level or, rather, on type II fiber recruitment. Thus, a change in the difference between Ceff and FCC can be explained by the change in the difference between Copt and Ceff. Other studies that have tried to explain the cadence paradox have shown a reduced difference between FCC and Ceff with fatigue.6,17,27 Argentin et al,27 for example, showed a significant shift in FCC (87–68 rpm) toward Ceff (65 rpm) after 2 hours of cycling at 65% of maximal aerobic power. As Copt decreases with fatigue, which is primarily from an increase in the internal power,14 the reduced difference between FCC and Ceff with fatigue can also be explained by the reduced difference between Copt and Ceff with fatigue. Regrettably, no study has simultaneously analyzed the influence of fatigue on FCC, Copt, and Ceff to confirm this theoretical assumption. The hypothesis that most competitive cyclists choose a cadence near Copt should be confirmed in future studies by simultaneously analyzing the influence of other “external” factors on FCC and Copt. The factors of interest could include Pext, fatigue, training, muscle-fiber-type composition, saddle height, and crank length. Furthermore, it is not assumed that all competitive cyclists chose an optimal cadence. Therefore, the percentage of cyclists that chose too high or too low a cadence must be estimated in a future study using a large number of subjects. Studies should also be performed with recreational cyclists and noncyclists to show if the subjects choose an optimal cadence independent of cycling experience. It is recommended that future studies be performed with more cadences tested to enhance precision in determining Copt. Level Ground Versus Uphill By comparing level-ground cycling with DP and uphill cycling with UP, this study had a second aim: to determine the influence of a concomitant change in road gradient and body position on FCC, Pmax, and Copt. Concerning FCC, experimental studies have revealed that it is influenced by road gradient3 but not by body position.28 Thus, a concomitant change in the road gradient and body position should theoretically have the same influence on FCC as only a change of road gradient. Our results on the treadmill support the results from Hansen et al,3 who also observed a lower FCC when subjects cycled uphill on a treadmill (69 rpm at 150 W and 73 rpm at 250 W) than while cycling on level ground (75 rpm at 150 W and 82 rpm at 250 W). These results also agree with observations of cyclists during their normal training29 or during competitions.30 A previous study showed that a change in the road gradient (0 vs 7%) alone influences Pmax and Copt and that a change in body position (DP vs UP) alone influences Pmax but not Copt.18 These results led to the assumption that Copt is lower and Pmax is higher for uphill cycling with UP than for level-ground cycling with DP. This assumption has now been experimentally confirmed. Practical Applications The results from this study suggest that most competitive cyclists freely choose a cadence near Copt to minimize fatigue at a given Pext. Thus, athletes should not attempt to copy the cadence used by successful cyclists and coaches should avoid forcing athletes to use a desired cadence. On the other hand, some cyclists freely choose too high or too low a cadence. Thus, the comparison of the individual FCC and the individual Copt at a specific endurance-performance level is an adequate method to optimize performance. The results from the second part of this study confirmed that under real cycling conditions it is advantageous from a performance-related point of view to use a lower cadence and a more upright body position during uphill cycling. Conclusion Despite the intense discussion on the FCC in cycling, the underlying factors that lead cyclists to choose a particular cadence remained under debate. This study demonstrated that interindividual, as well as intraindividual, differences in FCC are related to differences in Copt. These results indicate a causative relationship between FCC and Copt. As expected from a theoretical perspective, most cyclists freely choose a cadence near Copt to minimize peripheral fatigue at a given Pext or to maximize Pext for the given task. Thus, the cadence paradox, which states that competitive cyclists choose a higher cadence than Ceff, has been resolved. Furthermore, by comparing level-ground cycling with DP and uphill cycling with UP, this study confirmed that under real cycling conditions it is advantageous to use a lower cadence and a more upright body position during uphill cycling. Acknowledgments These studies were supported by the Swiss Federal Council of Sports (ESK/BASPO). The authors are grateful to the subjects for having participated in these studies. They also thank Marco Hitz and Peter Schwilch for their technical support. References 1. Vercruyssen F, Brisswalter J. Which factors determine the freely chosen cadence during submaximal cycling? J Sci Med Sport. 2010;13(2):225–231. PubMed doi:10.1016/j. jsams.2008.12.631 FCC and Optimal Cadence in Cycling 381 2.Hansen EA, Smith G. Factors affecting cadence choice during submaximal cycling and cadence influence on performance. Int J Sports Physiol Perform. 2009;4(1):3–17. PubMed 3. Hansen EA, Jorgensen LV, Jensen K, Fregly BJ, Sjogaard G. Crank inertial load affects freely chosen pedal rate during cycling. J Biomech. 2002;35(2):277–285 Erratum in J Biomech. 2002;35(2):1521. PubMed doi:10.1016/ S0021-9290(01)00182-8 4.Lucia A, Hoyos J, Chicharro JL. Preferred pedalling cadence in professional cycling. Med Sci Sports Exerc. 2001;33(8):1361–1366. PubMed 5. Hausswirth C, Lehenaff D, Dreano P, Savonen K. Effects of cycling alone or in a sheltered position on subsequent running performance during a triathlon. Med Sci Sports Exerc. 1999;31(4):599–604. PubMed doi:10.1097/00005768199904000-00018 6.Vercruyssen F, Hausswirth C, Smith D, Brisswalter J. Effect of exercise duration on optimal pedaling rate choice in triathletes. Can J Appl Physiol. 2001;26(1):44–54. PubMed 7. Hausswirth C, Argentin S, Bieuzen F, Le Meur Y, Couturier A, Brisswalter J. Endurance and strength training effects on physiological and muscular parameters during prolonged cycling. J Electromyogr Kinesiol. 2010;20(2):330–339. doi:10.1016/j.jelekin.2009.04.008. PubMed 8. Hansen EA, Andersen JL, Nielsen JS, Sjogaard G. Muscle fibre type, efficiency, and mechanical optima affect freely chosen pedal rate during cycling. Acta Physiol Scand. 2002;176(3):185–194. PubMed doi:10.1046/j.1365201X.2002.01032.x 9. Moussay S, Dosseville F, Gauthier A, Larue J, Sesboue B, Davenne D. Circadian rhythms during cycling exercise and finger-tapping task. Chronobiol Int. 2002;19(6):1137– 1149. PubMed doi:10.1081/CBI-120015966 10. Marais G, Pelayo P. Cadence and exercise: physiological and biomechanical determinants of optimal cadences— practical applications. Sports Biomech. 2003;2(1):103– 132. PubMed doi:10.1080/14763140308522811 11.MacIntosh BR, Neptune RR, Horton JF. Cadence, power, and muscle activation in cycle ergometry. Med Sci Sports Exerc. 2000;32(7):1281–1287. PubMed doi:10.1097/00005768-200007000-00015 12. Marsh AP, Martin PE, Sanderson DJ. Is a joint momentbased cost function associated with preferred cycling cadence? J Biomech. 2000;33(2):173–180. PubMed doi:10.1016/S0021-9290(99)00155-4 13. Takaishi T, Yasuda Y, Ono T, Moritani T. Optimal pedaling rate estimated from neuromuscular fatigue for cyclists. Med Sci Sports Exerc. 1996;28(12):1492–1497. PubMed 14.Emanuele U, Denoth J. Power–cadence relationship in endurance cycling. Eur J Appl Physiol. 2012;112(1):365– 375. PubMed doi:10.1007/s00421-011-1987-z 15. Coast JR, Welch HG. Linear increase in optimal pedal rate with increased power output in cycle ergometry. Eur J Appl Physiol Occup Physiol. 1985;53(4):339–342. PubMed doi:10.1007/BF00422850 16.Kohler G, Boutellier U. The generalized force–velocity relationship explains why the preferred pedaling rate of cyclists exceeds the most efficient one. Eur J Appl Physiol. 2005;94(1-2):188–195. PubMed doi:10.1007/s00421-0041283-2 17.Brisswalter J, Hausswirth C, Smith D, Vercruyssen F, Vallier JM. Energetically optimal cadence vs. freelychosen cadence during cycling: effect of exercise duration. Int J Sports Med. 2000;21(1):60–64. PubMed doi:10.1055/s-2000-8857 18. Emanuele U, Denoth J. Influence of road incline and body position on power–cadence relationship in endurance cycling. Eur J Appl Physiol. 2012;112(7):2433–2441. doi:10.1007/s00421-011-2213-8. PubMed 19. Baron B, Noakes TD, Dekerle J, et al. Why does exercise terminate at the maximal lactate steady state intensity? Br J Sports Med. 2008;42(10):828–833. PubMed doi:10.1136/ bjsm.2007.040444 20.Tegtbur U, Busse MW, Braumann KM. Estimation of an individual equilibrium between lactate production and catabolism during exercise. Med Sci Sports Exerc. 1993;25(5):620–627. PubMed 21. Fontana P, Boutellier U, Knopfli-Lenzin C. Time to exhaustion at maximal lactate steady state is similar for cycling and running in moderately trained subjects. Eur J Appl Physiol. 2009;107(2):187–192. PubMed doi:10.1007/ s00421-009-1111-9 22.Coleman DA, Wiles JD, Davison RC, Smith MF, Swaine IL. Power output measurement during treadmill cycling. Int J Sports Med. 2007;28(6):525–530. PubMed doi:10.1055/s-2006-955888 23. Marsh AP, Martin PE. The relationship between cadence and lower extremity EMG in cyclists and noncyclists. Med Sci Sports Exerc. 1995;27(2):217–225. PubMed 24. Nielsen JS, Hansen EA, Sjogaard G. Pedalling rate affects endurance performance during high-intensity cycling. Eur J Appl Physiol. 2004;92(1-2):114–120. PubMed doi:10.1007/s00421-004-1048-y 25. MacIntosh BR, Svedahl K, Kim M. Fatigue and optimal conditions for short-term work capacity. Eur J Appl Physiol. 2004;92(4-5):369–375. PubMed doi:10.1007/ s00421-004-1177-3 26. Foss O, Hallen J. The most economical cadence increases with increasing workload. Eur J Appl Physiol. 2004;92(45):443–451. PubMed doi:10.1007/s00421-004-1175-5 27. Argentin S, Hausswirth C, Bernard T, et al. Relation between preferred and optimal cadences during two hours of cycling in triathletes. Br J Sports Med. 2006;40(4):293–298, discussion 298. PubMed doi:10.1136/bjsm.2005.020487 28.Jobson SA, Nevill AM, George SR, Jeukendrup AE, Passfield L. Influence of body position when considering the ecological validity of laboratory time-trial cycling performance. J Sports Sci. 2008;26(12):1269–1278. PubMed doi:10.1080/02640410802183585 29. Sassi A, Rampinini E, Martin DT, Morelli A. Effects of gradient and speed on freely chosen cadence: the key role of crank inertial load. J Biomech. 2009;42(2):171–177. PubMed doi:10.1016/j.jbiomech.2008.10.008 30. Lucia A, Hoyos J, Chicharro JL. Physiology of professional road cycling. Sports Med. 2001;31(5):325–337. PubMed doi:10.2165/00007256-200131050-00004