International SportMed Journal
Optimal cadence selection during cycling
*1, 2, 3Dr Chris R Abbiss, PhD, 4Dr Jeremiah J Peiffer, PhD, 1Associate Professor Paul B Laursen, PhD
1School of Exercise, Biomedical and Health Sciences, Edith Cowan University, Joondalup, WA, Australia
2Department of Physiology, Australian Institute of Sport, Belconnen, ACT, Australia
3Division of Materials Science and Engineering, Commonwealth Scientific and Industrial Research Organisation, Belmont, Vic, Australia
4Centre of Excellence in Alzheimer’s Disease Research and Care, Vario Health Institute, Edith Cowan University, Joondalup, WA, Australia
Cadence or pedal rate is widely accepted as an important factor influencing economy of motion, power output, perceived exertion and the development of fatigue during cycling. As a result, the cadence selected by a cyclist’s could have a significant influence on their performance. Despite this, the cadence that optimises performance during an individual cycling task is currently unclear. The purpose of this review therefore was to examine the relevant literature surrounding cycling cadence in order provide a greater understanding of how different cadences might optimise cycling performance. Based on research to date, it would appear that relatively high pedal rates (100-120rpm) improve sprint cycling performance, since muscle force and neuromuscular fatigue are reduced, and cycling power output maximised at such pedal rates. However, extremely high cadences increase the metabolic cost of cycling. Therefore prolonged cycling (i.e. road time trials) may benefit from a slightly reduced cadence (~90-100rpm). During ultra-endurance cycling (i.e. >4h), performance might be improved through the use of a relatively low cadence (70-90rpm), since lower cadences have been shown to improve cycling economy and lower energy demands. However, such low cadences are known to increase the pedal forces necessary to maintain a given power output. Future research is needed to examine the multitude of factors known to influence optimal cycling cadence (i.e. economy, power output and fatigue development) in order to confirm the range of cadences that are optimal during specific cycling tasks. Keywords: pedal rate, economy, efficiency, power output
*Dr Chris R Abbiss, PhD
Dr Abbiss is a post-doctoral research fellow in high performance cycling at Edith Cowan University, Australian Institute of Sport and the Commonwealth Scientific and Industrial Research Organisation, Australia. His primary research interests centre on human physiology and exercise performance, with focuses on cycling, fatigue, thermoregulation, pacing strategies and training. His work in this area has resulted in the publication of numerous peer-reviewed scientific articles.
Dr Jeremiah Peiffer, PhD
Dr Peiffer is a post-doctoral research fellow at the Centre of Excellence for Alzheimer's Disease Research and Care in the School of Exercise, Biomedical and Health Sciences and Vario Health Institute at Edith Cowan Univ ersity in Perth, Australia. His research interests focus on the influence of physical activity on ageing, chronic disease, and Alzheimer's disease as well as cycling research focusing on thermoregulation, recovery, and fatigue.
Email address: email@example.com
Dr Laursen is an Associate Professor of Exercise Physiology in the School of Exercise, Biomedical and Health Sciences at Edith Cowan University, Perth Australia. His research interests centre on various aspects of human physiology, with focuses on fatigue, high-intensity training, thermoregulation, hydration, muscle damage and recovery kinetics.
Email address: firstname.lastname@example.org
Understanding factors that affect cycling performance is of interest to scientists, coaches and cyclists alike. Accordingly, a vast body of literature has examined how various environmental, physiological and biomechanical factors influence cycling performance (for review, see references 23, 24). From this work, cycling performance would appear to be dictated largely by the ability of the cyclist to produce high power outputs at minimal metabolic costs. As pedal rate (i.e. cadence) can influence both the ability to produce power, as well as rate of energy consumption, cadence selection could have a significant impact on cycling performance. For instance, the adoption of a high cadence (~90rpm) has been shown to reduce myoelectrical activity, muscle force and neuromuscular fatigue 64. In contrast, high cadences (80-120rpm) have also been found to be less economical than lower cadences (~90rpm) 15. Indeed, Bieuzen et al. 9 observed a difference between energetically optimal cadence and neuromuscular optimal cadence (63.5 and 93.5rpm, respectively), in well trained cyclists. In addition, optimal and self-selected cadences have been found to be influenced by cycling intensity 46, course geography (grade) 43, muscle fibre composition 18, 33, 52 and cycling experience 46. While information concerning pedal rate selection during cycling exists, a comprehensive review of the present literature is not currently available. As such, the cadence that results in the best possible performance outcome during the vast array of cycling events and conditions remains unclear. Therefore the purpose of this review is to 1) examine the literature pertaining to self-selected, forced and optimal cadences, 2) determine the factors that are responsible for the self-selection of cadence, and 3) provide a greater understanding of cadences that optimise performance during the variety of tasks performed by cyclists.
Understanding self-selected/freely chosen cadence
Publications on cycling cadence often comment on the unusually high pedal rate (>90rpm) adopted during both level and uphill cycling by seven time Tour de France champion Lance Armstrong 17, 41. Based on the success of this cyclist it seems reasonable to propose that such high pedal rates might optimise performance during the most influential phases of professional cycling events (i.e. uphill and time trials). However, successful elite cyclists have also been observed to adopt significantly lower cadences during uphill mountain accents (>90rpm) 43, 74. As such, the effect of such high cadences on performance during cycling is unclear. It has been suggested that a high mechanical efficiency and maximal aerobic capacity (V•O2max) may allow particular cyclists to increase power output (475 – 500W) using noticeably higher pedal rates 42. Alternatively, other cyclists may choose to cycle at lower cadences in order to minimise oxygen cost, since higher cadences appear to be less efficient (see section on Efficiency and economy). If this hypothesis is correct, then any one single cycling cadence is not likely to be beneficial for all cyclists 42. Instead, the optimal cadence to adopt during cycling will depend on the central (i.e. V•O2max) and peripheral (i.e. muscle fibre contribution) physiological characteristics of each individual. Such a notion could explain the close association observed between an athlete’s cycling experience and a freely chosen pedal rate. Indeed, well-trained cyclists typically adopt higher cadences compared with their lesser trained counterparts 16, 29, 46. However, as with optimal cadences, the factors affecting self-selected pedal rate remain unclear.
Defining optimal cadence
For many years, scientists, coaches and athletes have attempted to determine the optimal pedal rate to apply during a variety of cycling tasks. While numerous investigations have been conducted 25, 28, 59, 66, 69, the best possible cycling cadence remains unclear. This uncertainty may be due to methodological differences and variations in the precise definition of the term ‘optimal’ used within cadence research. Indeed, previous research in this area has focused on the effects of various cadences on cycling mechanics 38, 49, 66, cycling efficiency 44, hemodynamics 28, 69, neuromuscular fatigue 37, 64, 65, 72 and more recently cycling performance 25, 75. Therefore the ideal cycling cadence may differ, dependent on whether the term refers to the most economical, powerful, fatigue-resistant or comfortable cadence 25, 57. For the purpose of this review on cadence, the term ‘optimal’ refers to the pedal rate resulting in the best possible performance outcome. The cadence that optimises performance under a variety of conditions experienced by cyclists is likely to be dictated by the trade-off between cycling economy, power output and the development of fatigue. Thus throughout this review each of these variables will be discussed with respect to various cycling disciplines.
Cycling power output
Pedal force and joint moments
Recent advancements in strain-gauge technology have led to improved understanding of the interaction between pedal force and resultant crank torque 7, 38, 49, 57. Studies examining pedal force during cycling have revealed both effective and ineffective pedal loads 11, 22, 61. Further, effective and ineffective pedal loads can be separated, by force, into normal (force applied perpendicularly to the pedal surface) and tangential (force applied along the surface of the pedal) components. Typical effective pedal loading at various cadences and power outputs are shown in Figure 1.
Figure 1: Effective pedal forces throughout the entire crank cycle during cycling at various power outputs (a) and cadences (b). 0° crank angle refers to top dead centre. Figure reproduced by permission of the publisher Taylor & Francis Ltd , http://www.tandf.co.uk/journals from Sanderson DJ, Hennig EM and Black AH. The influence of cadence and power output on force application and in-shoe pressure distribution during cycling by competitive and recreational cyclists. J of Sports Sci 2000. 61.
The effective pedal force acts perpendicularly to the bicycle crank, generating a torque which is transmitted through the bicycle chain to the wheel. From Figure 1 it can be seen that the effective pedal force and thus crank torque vary substantially throughout the pedal cycle. Indeed, peak torque typically occurs at approximately 100°, past top dead centre, whereas negative load occurs on the upstroke of the pedal cycle (Figure 1). Since power output is the product of crank torque and crank angular velocity, instantaneous power output also varies throughout a crank cycle 11.
Despite this, average power output produced over an entire revolution may be determined by the following equation:
Power (W) = average net effective pedal torque x average angular velocity (cadence) 11.
Based on the above formula, the average force/torque applied to the pedals over an entire pedal revolution at a fixed power output is reduced at higher cadences 71. For example, at 350W the average effective force applied to the pedals is ~15% lower when cycling at 105rpm (184N) compared with 90rpm (215N). This reduction in average pedal force (i.e. force over an entire revolution) is predominantely due to a decrease in peak normal forces (Figure 1b; 60, 61). This is important as the peak normal forces observed during maximal cycling are likely to be dictated by the force/torque-velocity relationship of muscle contractions 45, 63. In short, the peak torque that can be applied to the pedals during short duration maximal cycling is reduced at faster contraction velocities (i.e. faster cadences; Figure 2).
Figure 2: Relationship between peak crank torque, crank velocity (i.e. cadence) and power output during short duration (<10s) maximal cycling in two separate subjects (solid and dashed lines). Figure used with permission from J Appl Physiol. 51
Based on the contractile properties of human muscle it has been shown that maximal cycling power output is achieved at approximately 120-130rpm (Figure 2; 8, 50, 51, 62, 78). Such high cadences may be important to maximal sprint cycling performance. Indeed, track and bicycle motorcross (BMX) cyclists typically perform short duration events (?m) at average cadences equal to or greater than 120 rpm 19, 20. However, Zoladz et al. 78 found that when pedalling above 100rpm there was a decrease in the power output delivered at any given oxygen cost, which was in turn associated with an earlier onset of anaerobiosis 77, 78. Such findings highlight the disadvantage of adopting such high cadences (>100rpm) during prolonged high-intensity exercise, such as competitive road cycling.
Inertial load and momentum
To further understand the mechanical cost associated with cadence, researchers have quantified both the muscular and non-muscular components that dictate pedal forces and crank torques 7, 54. Muscular components refer to forces or torques that are produced by muscular activity, whereas non-muscular components refer to other factors that may influence pedal or crank forces, such as gravity or inertia 7. While muscular pedal forces are reduced with higher cadences (as described in the previous section), non-muscular pedal forces increase linearly with pedal rate 7, 54. As a result, overall pedal forces at fixed power outputs follow a quadratic relationship with increases in cadence (Figure 3).
Figure 3: Muscular, non-muscular and total pedal forces while cycling at 120W and at 60, 75, 90, 105 and 120rpm. Reprinted from: J Biomech, Vol.32, Neptune RR and Hertzog W. The association between negative muscle work and pedaling rate, pp.1051-1026, 1999, with permission from Elsevier. 54.
Examining this relationship, Neptune and Herzog 54 found that when cycling at 260W, a minimum pedal force of 190N was observed at 90rpm, compared with higher (105 and 120rpm) and lower (60 and 75rpm) cadences. Since gravitational forces are largely unaffected by changes in cadence 7, 12, increasing non-muscular pedal forces that occur with higher cadences are primarily due to the influence of inertial load (i.e. increased inertia of the crank) (kg m2) 7. Indeed, by increasing crank inertial loads, self-selected cadence has been found to significantly increase, possibly an attempt to reduce peak crank torque 31. It has therefore been suggested that changes in inertial properties associated with increasing cadence may influence lower extremity neuromuscular coordination 7, 38, 40.
Neuromuscular fatigue and myoelectrical activity
The influence of inertial properties on neuromuscular coordination during cycling has previously been examined 7, 40, 55. Through the examination of muscle activation burst patterns and the coordination of mono- and bi-articular antagonists, it has been shown that higher cadences result in a forward shift (i.e. earlier in the crank cycle) in the activation of gluteus maximus, vastus lateralis, and tibialis anterior 38, 39. Further, the magnitude of this shift decreased in proximal (hip) to distal (ankle) limb segments 39. Since increases in pedalling rate can influence neuromuscular recruitment patterns, it seems plausible to presume that variations in cycling cadence might induce the development of neuromuscular fatigue. Nevertheless, research on this topic has produced conflicting results 37, 44, 64-66, 72. Studies have shown that the self-selection of relatively high pedalling rates (~80-90rpm) may reduce muscle activation of vastus lateralis 44, 64, 72. Therefore it has been suggested that cyclists may spontaneously select a high cadence in order to prevent the development of neuromuscular fatigue, regardless of the energy cost 64. In support of this, Takaishi et al. 72 found that integrated electromyography (iEMG) of vastus lateralis as a function of time (i.e. slope of the iEMG) followed a quadratic relationship with cadence and was minimised at 80-90rpm. The authors concluded that cyclists tend to adopt such cadences in order to minimise muscular fatigue, and not metabolic demand, since lower cadences (60-70rpm) were associated with reduced oxygen consumption 71, 72. In contrast to these findings, Sarre et al. 65 found that when cycling at constant power outputs (60%, 80% and 100% of maximal aerobic power output) neuromuscular activation of vastus lateralis and vastus medialis were unaffected by varying cadence (70 – 130% of self-selected pedal rate). Further, with the use of femoral nerve electrical stimulation, it was shown that similar variations in cadence (± 20% of self-selected cadence) during prolonged cycling had no effect on the occurrence of either central or peripheral fatigue development of the leg extensors 37, 66. Inconsistencies in findings within this research area may be related to methodological differences relating to the functional role of the muscles examined, training level of the subjects, and the range of power outputs/cadences used 64.
Figure 4. Diagram demonstrating the direction and magnitude of pedal forces throughout a typical clockwise pedal cycle. Note counterproductive (negative) pedal forces during the upward phase of the pedal cycle. Reprinted, with permission, from J.P. Broker, 2003, Cycling Biomechanics: Road and Mountain. In: High-Tech Cycling: The Science of Riding Faster, edited by E.R. Burke, 2nd ed. (Champaign, IL: Human Kinetics, 125, figure 5.4.11.
This negative force is generated by the insufficient speed of the rear leg during the upstroke of the pedal cycle 57, 66. Despite an increase in activation of biceps femoris at higher cadences, the increase in pedal rate and thus crank speed/momentum may still result in greater negative work. Thus in order to overcome this increase in negative work, it has been suggested that the front leg may be required to perform greater positive work during the downstroke, resulting in increased fatigue (especially at high power outputs) 66.
Efficiency and economy
Numerous studies have examined the influence of pedalling frequency on the efficiency and economy of cycling 6, 27, 44, 46, 48, 71, 72. Generally, when cycling at constant power outputs, lower cadences have been found to result in reduced oxygen cost (i.e. improved gross efficiency) compared with higher cadences 6, 15, 27. Improved efficiency of cycling observed at lower pedalling rates is likely to be dictated by the relationship between muscle shortening velocity and the efficiency of muscle contractions (percent Type I and Type II active fibres). For instance, under in vitro conditions, it has been observed that the efficiency of skeletal muscle contractions is augmented with increasing speed of contraction, until a maximum is reached (i.e. an economically optimal shortening velocity) 35. The most economical cadence appears to be extremely low (~50-60rpm) when cycling at low power outputs (?W), but increases to approximately 80-100rpm with increasing workloads (~350W) 26, 44, 58. The cause of the rise in the economically optimal cadence is unclear, but is again likely to be due to the power-velocity relationship of muscle contraction and the additional recruitment of fast twitch muscle fibres with increases in exercise intensity. As previously mentioned (see section on Pedal force and joint moments), an increase in cadence at higher exercise intensities may optimise the power-velocity relationship, and as a result reduce the metabolic cost of cycling. Indeed at lower cadences, greater force per pedal stroke is required to maintain a given power output, which requires additional muscle fibre recruitment and thus a higher energy expenditure 58, 67. Supporting this, myoelectrical activity of vastus lateralis is reduced at higher cadences (see section on Neuromuscular fatigue and myoelectrical activity).
While numerous studies have investigated the influence of cadence on oxygen consumption, power output and fatigue development (described above), few studies have examined the hemodynamic changes associated with different pedal rates 13, 28, 67, 69, 70. As with oxygen consumption, it has been observed that when cycling at a constant power output, heart rate may increase both above and below the ideal cadence 67. Further, the pedal rate that minimises heart rate rises linearly with increasing power output 13, 67. The close relationship between the energetically optimal cadence (i.e. cadence which minimises V•O2) and the cadence which minimises heart rate may be related to the oxygen (see section on Efficiency and economy) and thus blood flow demands of working muscle. Gotshall et al. 28 have shown increases in heart rate, stroke volume and cardiac output with higher pedal cadences ranging from 70-110rpm. However, in this study the elevated cardiac output observed at higher cadences was associated with a disproportionately lower rise in oxygen consumption, as shown by a reduction in the arterial-venous oxygen difference 28. Consequently, this study showed that increases in cardiac output observed at higher cadences were not solely due to elevated oxygen demands. Instead the authors suggested that the higher cardiac output could have been due to the enhanced effectiveness of the skeletal muscle pump resulting from the faster cadences 28. Indeed, the greater contraction rate occurring at higher cadences would facilitate venous return, augment ventricular preload, and elevate cardiac output.
In addition to increasing venous return, higher cadences might also reduce the period of blood flow occlusion that occurs in the microvessels of skeletal muscle during cycling. With the use of near infrared spectroscopy (NIRS), Takaishi and co-workers 69, 70 found that when cycling at 75% V•O2max, muscle blood flow and oxygenation of vastus lateralis was significantly reduced during the initial pedal downstroke (first third of the crank cycle; Figure 5), presumably due to high intramuscular pressure associated with muscle contraction.
Figure 5: Changes in muscle blood flow (circle), muscle oxygenation (square), pedal forces (triangle), knee angle and rectified electromyography (EMG) throughout a pedal cycle. 0° crank angle refers to top dead centre. Figure reprinted with permission from Lippincott Williams & Wilkens for Takaishi T, Sugiura T, Katayama K, et al. Changes in blood volume and oxygenation level in a working muscle during a crank cycle, Med Sci Sports Exerc, Vol.34 No.3, 2002, pp.520-528 70.
Further, in untrained individuals, this deoxygenation (i.e. the minimum blood volume and oxygenation reached) was more sever at low (50rpm) compared with high (85rpm) cadences 69. It is therefore plausible that higher cadences could improve oxygen delivery to working muscles by limiting blood flow occlusion. Such findings may be especially important during the forceful contractions (i.e. during high power outputs) typically achieved by professional/elite cyclists. Despite this, further research is needed in order to determine the influence of hemodynamics on preferred and optimal cadence selection during cycling.
A vast body of literature has examined various factors that may influence the optimal pedal rate to adopt during a variety of cycling tasks. Despite this research, the cadence which maximises performance during cycling remains unclear. It is possible that much of the uncertainty surrounding optimal cadences could be due to methodological inconsistencies between studies. In particular, the term ‘optimal’ may be used to describe the most economical, powerful, fatigue-resisting or comfortable pedal rates. As a result, the cadence that results in the best possible performance during the variety of cycling tasks experienced by cyclists appears to be multifaceted. Consequently, future research exploring the best possible cadence to select during cycling should examine a number of factors (i.e. power, neuromuscular fatigue, efficiency, blood flow and comfort) that may be associated with maximising performance outcomes. In particular, the influence of training at various cadences on performance and physiological adaptations requires further examination. Based on previous research, it would appear that muscle force and neuromuscular fatigue might be reduced, and cycling power output maximised, with relatively high pedal rates (100-120rpm). However, such high pedal rates increase the metabolic cost of cycling, especially at low power outputs (≤ 200W). As a result, short duration sprint cycling performance might be optimised with the adoption of fast pedal rates (~120rpm). Due to the influence that fast pedal rates have been shown to impart on cycling mechanics, cycling efficiency and fatigue development, performance in longer duration events might be enhanced from use of slightly slower cadences (~90-100rpm). During ultra-endurance cycling, performance might be improved by using relatively low cadences (70-90rpm), since cycling economy is improved and energy demands are lowered. Future research examining a multitude of factors known to influence optimal cycling cadence (i.e. economy, power output and fatigue development) is needed to confirm these hypotheses.
Dr Chris R Abbiss, School of Exercise, Biomedical and Health Sciences, Edith Cowan University, 100 Joondalup Drive, Joondalup, WA, Australia 6027
Tel.: +61 8 6304 5740
Fax: + 61 8 6304 5036
1. Abbiss CR, Quod MJ, Martin DT, et al. Dynamic pacing strategies during the cycle phase of an Ironman triathlon. Med Sci Sports Exerc 2006; 38: 726-734.
2. Ahlquist LE, Bassett DR, Jr., Sufit R, et al. The effect of pedaling frequency on glycogen depletion rates in type I and type II quadriceps muscle fibers during submaximal cycling exercise. Eur J Appl Physiol Occup Physiol 1992; 65: 360-364.
3. Arendt-Nielsen L, and Mills KR. The relationship between mean power frequency of the EMG spectrum and muscle fibre conduction velocity. Electroencephalogr Clin Neurophysiol 1985; 60: 130-134.
4. Arendt-Nielsen L, Mills KR, and Forster A. Changes in muscle fiber conduction velocity, mean power frequency, and mean EMG voltage during prolonged submaximal contractions. Muscle Nerve 1989; 12: 493-497.
5. 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: 293-298; discussion 298.
6. Banister EW, and Jackson RC. The effect of speed and load changes on oxygen intake for equivalent power outputs during bicycle ergometry. Int Z Angew Physiol 1967; 24: 284-290.
7. Baum BS, and Li L. Lower extremity muscle activities during cycling are influenced by load and frequency. J Electromyogr Kinesiol 2003; 13: 181-190.
8. Beelen A, and Sargeant AJ. Effect of fatigue on maximal power output at different contraction velocities in humans. J Appl Physiol 1991; 71: 2332-2337.
9. Bieuzen F, Vercruyssen F, Hausswirth C, et al. Relationship between strength level and pedal rate. Int J Sports Med 2007; 28: 585-589.
10. Brisswalter J, Hausswirth C, Smith D, et al. Energetically optimal cadence vs. freely-chosen cadence during cycling: effect of exercise duration. Int J Sports Med 2000; 21: 60-64.
11. Broker JP. Cycling biomechanics: Road and mountain. In: Burke ER, ed. High-Tech Cycling. Champaign: Human Kinetics, 2003.
12. Brown DA, Kautz SA, and Dairaghi CA. Muscle activity patterns altered during pedaling at different body orientations. J Biomech 1996; 29: 1349-1356.
13. Candau RB, Grappe F, Menard M, et al. Simplified deceleration method for assessment of resistive forces in cycling. Med Sci Sports Exerc 1999; 31: 1441-1447.
14. Chapman AE, and Sanderson DJ. Muscular co-ordination in sporting skills. In: Winters JM, and Woo SL-Y, ed. Multiple Muscle Systems: Biomechanics and Movement Organization. New York: Springer-Verlag, 1990, p. 608-620.
15. Chavarren J, and Calbet JA. Cycling efficiency and pedalling frequency in road cyclists. Eur J Appl Physiol Occup Physiol 1999; 80: 555-563.
16. Coast JR, and Welch HG. Linear increase in optimal pedal rate with increased power output in cycle ergometry. Eur J Appl Physiol Occup Physiol 1985; 53: 339-342.
17. Coyle EF. Improved muscular efficiency displayed as Tour de France champion matures. J Appl Physiol 2005; 98: 2191-2196.
18. Coyle EF, Sidossis LS, Horowitz JF, et al. Cycling efficiency is related to the percentage of type I muscle fibers. Med Sci Sports Exerc 1992; 24: 782-788.
19. Craig NP, and Norton KI. Characteristics of track cycling. Sports Med 2001; 31: 457-468.
20. Dorel S, Hautier CA, Rambaud O, et al. Torque and power-velocity relationships in cycling: relevance to track sprint performance in world-class cyclists. Int J Sports Med 2005; 26: 739-746.
21. Ericson MO. Mechanical muscular power output and work during ergometer cycling at different work loads and speeds. Eur J Appl Physiol Occup Physiol 1988; 57: 382-387.
22. Ericson MO, and Nisell R. Efficiency of pedal forces during ergometer cycling. Int J Sports Med 1988; 9: 118-122.
23. Faria EW, Parker DL, and Faria IE. The science of cycling: factors affecting performance - part 2. Sports Med 2005; 35: 313-337.
24. Faria EW, Parker DL, and Faria IE. The science of cycling: physiology and training - part 1. Sports Med 2005; 35: 285-312.
25. Foss O, and Hallen J. Cadence and performance in elite cyclists. Eur J Appl Physiol 2005; 93: 453-462.
26. Foss O, and Hallen J. The most economical cadence increases with increasing workload. Eur J Appl Physiol 2004; 92: 443-451.
27. Gaesser GA, and Brooks GA. Muscular efficiency during steady-rate exercise: effects of speed and work rate. J Appl Physiol 1975; 38: 1132-1139.
28. Gotshall RW, Bauer TA, and Fahrner SL. Cycling cadence alters exercise hemodynamics. Inter J Sports Med 1996; 17: 17-21.
29. Hagberg JM, Mullin JP, Giese MD, et al. Effect of pedaling rate on submaximal exercise responses of competitive cyclists. J Appl Physiol 1981; 51: 447-451.
30. Hagg GM. Interpretation of EMG spectral alterations and alteration indexes at sustained contraction. J Appl Physiol 1992; 73: 1211-1217.
31. Hansen EA, Jorgensen LV, Jensen K, et al. Crank inertial load affects freely chosen pedal rate during cycling. J Biomech 2002; 35: 277-285.
32. Hansen EA, Raastad T, and Hallen J. Strength training reduces freely chosen pedal rate during submaximal cycling. Eur J Appl Physiol 2007; 101: 419-426.
33. Hansen EA, and Sjogaard G. Relationship between efficiency and pedal rate in cycling: significance of internal power and muscle fiber type composition. Scand J Med Sci Sports 2007; 17: 408-414.
34. He ZH, Bottinelli R, Pellegrino MA, et al. ATP consumption and efficiency of human single muscle fibers with different myosin isoform composition. Biophys J 2000; 79: 945-961.
35. Heglund NC, and Cavagna GA. Mechanical work, oxygen consumption, and efficiency in isolated frog and rat muscle. Am J Physiol 1987; 253: C22-29.
36. Hull ML, and Gonzalez H. Bivariate optimization of pedalling rate and crank arm length in cycling. J Biomech 1988; 21: 839-849.
37. Lepers R, Millet GY, and Maffiuletti NA. Effect of cycling cadence on contractile and neural properties of knee extensors. Med Sci Sports Exerc 2001; 33: 1882-1888.
38. Li L. Neuromuscular control and coordination during cycling. Res Q Exerc Sport 2004; 75: 16-22.
39. Li L, and Caldwell GE. Coefficient of cross correlation and the time domain correspondence. J Electromyogr Kinesiol 1999; 9: 385-389.
40. Li L, and Caldwell GE. Effect of inertial loading in muscle activity in cycling. Proceedings of ISBS XI, University of Massachusetts 1993; 120-125.
41. Lucia A, Earnest C, and Arribas C. The Tour de France: a physiological review. Scand J Med Sci Sports 2003; 13: 275-283.
42. Lucia A, Earnest C, Hoyos J, et al. Optimizing the crank cycle and pedaling cadence. In: Burke ER, ed. High-Tech Cycling. Champaign: Human Kinetics, 2003.
43. Lucia A, Hoyos J, and Chicharro JL. Preferred pedalling cadence in professional cycling. Med Sci Sports Exerc 2001; 33: 1361-1366.
44. Lucia A, San Juan AF, Montilla M, et al. In professional road cyclists, low pedaling cadences are less efficient. Med Sci Sports Exerc 2004; 36: 1048-1054.
45. MacIntosh BR, Neptune RR, and Horton JF. Cadence, power, and muscle activation in cycle ergometry. Med Sci Sports Exerc 2000; 32: 1281-1287.
46. Marsh AP, and Martin PE. Effect of cycling experience, aerobic power, and power output on preferred and most economical cycling cadences. Med Sci Sports Exerc 1997; 29: 1225-1232.
47. Marsh AP, and Martin PE. The relationship between cadence and lower extremity EMG in cyclists and noncyclists. Med Sci Sports Exerc 1995; 27: 217-225.
48. Marsh AP, Martin PE, and Foley KO. Effect of cadence, cycling experience, and aerobic power on delta efficiency during cycling. Med Sci Sports Exerc 2000; 32: 1630-1634.
49. Marsh AP, Martin PE, and Sanderson DJ. Is a joint moment-based cost function associated with preferred cycling cadence? J Biomech 2000; 33: 173-180.
50. McCartney N, Heigenhauser GJ, Sargeant AJ, et al. A constant-velocity cycle ergometer for the study of dynamic muscle function. J Appl Physiol 1983; 55: 212-217.
51. McCartney N, Obminski G, and Heigenhauser GJ. Torque-velocity relationship in isokinetic cycling exercise. J Appl Physiol 1985; 58: 1459-1462.
52. Mogensen M, Bagger M, Pedersen PK, et al. Cycling efficiency in humans is related to low UCP3 content and to type I fibres but not to mitochondrial efficiency. J Physiol 2006; 571: 669-681.
53. Mornieux G, Guenette JA, Sheel AW, et al. Influence of cadence, power output and hypoxia on the joint moment distribution during cycling. Eur J Appl Physiol 2007; 102: 11-18.
54. Neptune RR, and Herzog W. The association between negative muscle work and pedaling rate. J Biomech 1999; 32: 1021-1026.
55. Neptune RR, Kautz SA, and Hull ML. The effect of pedaling rate on coordination in cycling. J Biomech 1997; 30: 1051-1058.
56. Pandolf KB, and Noble BJ. The effect of pedalling speed and resistance changes on perceived exertion for equivalent power outputs on the bicycle ergometer. Med Sci Sports 1973; 5: 132-136.
57. Patterson RP, and Moreno MI. Bicycle pedalling forces as a function of pedalling rate and power output. Med Sci Sports Exerc 1990; 22: 512-516.
58. Pierre S, Nicolas H, and Fre´de´rique H. Interactions between cadence and power output effects on mechanical efficiency during sub maximal cycling exercises. Eur J Appl Physiol 2006; 97: 133-139.
59. Redfield R, and Hull ML. On the relation between joint moments and pedalling rates at constant power in bicycling. J Biomech 1986; 19: 317-329.
60. Sanderson DJ. The influence of cadence and power output on the biomechanics of force application during steady-rate cycling in competitive and recreational cyclists. J Sports Sci 1991; 9: 191-203.
61. Sanderson DJ, Hennig EM, and Black AH. The influence of cadence and power output on force application and in-shoe pressure distribution during cycling by competitive and recreational cyclists. J Sports Sci 2000; 18: 173-181.
62. Sargeant AJ. Effect of muscle temperature on leg extension force and short-term power output in humans. Eur J Appl Physiol Occup Physiol 1987; 56: 693-698.
63. Sargeant AJ. Human power output and muscle fatigue. Int J Sports Med 1994; 15: 116-121.
64. Sarre G, and Lepers R. Neuromuscular function during prolonged pedalling exercise at different cadences. Acta Physiol Scand 2005; 185: 321-328.
65. Sarre G, Lepers R, Maffiuletti N, et al. Influence of cycling cadence on neuromuscular activity of the knee extensors in humans. Eur J Appl Physiol 2003; 88: 476-479.
66. Sarre G, Lepers R, and van Hoecke J. Stability of pedalling mechanics during a prolonged cycling exercise performed at different cadences. J Sports Sci 2005; 23: 693-701.
67. Seabury JJ, Adams WC, and Ramey MR. Influence of pedalling rate and power output on energy expenditure during bicycle ergometry. Ergonomics 1977; 20: 491-498.
68. Suzuki Y. Mechanical efficiency of fast- and slow-twitch muscle fibers in man during cycling. J Appl Physiol 1979; 47: 263-267.
69. Takaishi T, Ishida K, Katayama K, et al. Effect of cycling experience and pedal cadence on the near-infrared spectroscopy parameters. Med Sci Sports Exerc 2002; 34: 2062-2071.
70. Takaishi T, Sugiura T, Katayama K, et al. Changes in blood volume and oxygenation level in a working muscle during a crank cycle. Med Sci Sports Exerc 2002; 34: 520-528.
71. Takaishi T, Yamamoto T, Ono T, et al. Neuromuscular, metabolic, and kinetic adaptations for skilled pedaling performance in cyclists. Med Sci Sports Exerc 1998; 30: 442-449.
72. Takaishi T, Yasuda Y, Ono T, et al. Optimal pedaling rate estimated from neuromuscular fatigue for cyclists. Med Sci Sports Exerc 1996; 28: 1492-1497.
73. Vercruyssen F, Hausswirth C, Smith D, et al. Effect of exercise duration on optimal pedaling rate choice in triathletes. Can J Appl Physiol 2001; 26: 44-54.
74. Vogt S, Roecker K, Schumacher YO, et al. Cadence-power-relationship during decisive mountain ascents at the Tour de France. Int J Sports Med 2008; 29: 244-250.
75. Watson G, and Swensen T. Effects of altering pedal cadence on cycling time-trial performance. Int J Sports Med 2006; 27: 296-300.
76. Woledge RC. Possible effects of fatigue on muscle efficiency. Acta Physiol Scand 1998; 162: 267-273.
77. Woolford SM, Withers RT, Craig NP, et al. Effect of pedal cadence on the accumulated oxygen deficit, maximal aerobic power and blood lactate transition thresholds of high-performance junior endurance cyclists. Eur J Appl Physiol Occup Physiol 1999; 80: 285-291.
78. Zoladz JA, Rademaker AC, and Sargeant AJ. Human muscle power generating capability during cycling at different pedalling rates. Exp Physiol 2000; 85: 117-124.
* Corresponding author. Address at the end of text.