Journal of Strength and Conditioning Research, 2006, 20(3), 547-554© 2006 National Strength & Conditioning Association

ASSISTED AND RESISTED SPRINT TRAININGIN SWIMMING

SEBASTIEN GIROLD,' PAUL CALMELS,^ DIDIER MAURIN/ NICOLAS MILHAU,' ANDJEAN-CLAUDE CHATARD'

'Laboratory of Physiology, PPEH, Faculty of Medicine Jean Monnet, Saint-Etienne, France; '^Department ofFunctional Rehabilitation, Hospital of St-Etienne, France,

ABSTRACT. Girold, S., P. Calmels, D. Maurin, N. Milhau, andJ.-C. Chatard. Assisted and resisted sprint training in swim-ming. J. Strength Cond. Res. 20(3}-.547-554. 2006.—This studywas undertaken to determine whether the resisted-sprint in ov-erstrength (OSt) or the assisted-sprint in overspeed (OSp) couldbe efficient training methods to increase 100-m front crawl per-formance. Thirty-seven (16 men, 21 women) competition-levelswimmers (mean ± SD\ age 17.5 ± 3.5 years, height 173 ± 14cm, weight 63 ± 14 kg) were randomly divided into 3 groups:OSt, OSp, and control (C). All swimmers trained 6 days per weekfor 3 weeks, including 3 resisted or assisted training sessionsper week for the groups OSt and OSp respectively. Elastic tubeswere used to generate swimming overstrength and overspeed.Three 100-m events were performed before, during, and afterthe training period. Before each 100-m event, strength of theelbow flexors and extensors was measured with an isokineticdynamometer. Stroke rate and stroke length were evaluated us-ing the video-recorded 100-m events. In the OSt group, elbowextensor strength, swimming velocity, and stroke rate signifi-cantly increased (p < 0.05), while stroke length remained un-changed after the 3-week training period. In the OSp group,stroke rate significantly increased (p < 0.05) and stroke lengthsignificantly decreased (p < 0.05) without changes in swimmingvelocity. No significant variations in the C group were observed.Botb OSt and OSp proved to be more efficient than the tradi-tional training program. However, the OSt training programbad a larger impact on muscle strength, swimming performance,and stroke technique than the OSp program.

KEY WORDS, exercise, muscular strength, overspeed, over-strength, performance

INTRODUCTION

trength and speed are two major elements de-termining a sprint swimmer's train^ing pro-gram, and physiological and technical param-eters are known to affect sprint swimmingperformance. Physiological parameters main-

ly concern strength and power of the upper limhs (11, 12,21). The main technical parameters are stroke rate anddistance per stroke or stroke length (1, 13, 15, 24). Insprint swimming, physical strength is considered moreimportant than technique to reach a high level of perfor-mance, although some variations have been observedwith gender and morphological parameters (13).

Several training methods have been described to de-velop strength in swimming (6, 10, 19, 28). Their efficien-cy depends on the specificity of the event (7, 25) and theintensity of the training sessions (2, 5, 6, 15, 18, 26).

The effects of high-resistance (strength) and high-ve-locity (speed) training programs on sprint performancehave been studied but only in running (8, 9). The resultsof these studies showed that the high-resistance training

method improves muscular strength without improving100-m performances, while the high-velocity trainingmethod increased 100-m performances without increas-ing muscular strength. This tj^je of study has not beenperformed with swimmers.

The main purpose of the present study was to deter-mine whether strength and performance can be developedin swimmers using a resisted-sprint (high-resistance) andan assisted-sprint (high-velocity) training program. A sec-ond purpose was to determine whether there is a rela-tionship between development of strength, technical pa-rameters, and performance in the different training pro-grams.

METHODS

Experimental Approach to the Prohiem

The main purpose of this study was to increase the per-formances of swimmers over 100 m using resisted- andassisted-sprint training methods. It was thus decided toperform these conditions while swimming using specificelastics in resisted (high-resistance) and assisted (high-velocity) conditions.

Specific sets were developed to develop sprint abilitiesover 100 m (with an anaerobic capacity dominant work).For practical reasons concerning the swimmers andcoaches, it was impossible to keep the training conditionsfor more than 3 weeks for the whole group of swimmers.However, it was considered that adaptations could appearconsidering results of other studies. Indeed, Pichon et al.(19) and Strass et al. (26) reported adaptations aftertraining programs that lasted only 3 weeks.

Subjects

A group of 45 competitive (19 men, 26 women) swimmers(mean ± SD: age 16.5 ± 3 years, height 171 ± 13 cm,weight 67 ± 21 kg), from regional to national level, v fasstudied. An institutional review board approval for theproject was obtained from the University Committee onHuman Research. The swimmers under age of 18 andtheir parents signed an informed consent form, and thesubjects participated in the study on a voluntary basis.Svidmmers trained an average 12 ± 1.5 hours per week.All swimmers were sprinters, 100-m, or 200-m front crawlspecialists. Due to injuries or missed training sessions,only 37 swimmers (16 men, 21 women, age 17.5 ± 3.5years, height 173 ± 14 cm. weight 63 ± 14 kg) completedthe whole program, inducing some heterogeneity in themain characteristics of the 3 groups defined below.

547

548 GiROLD, CALMEUS, MAURIN ET AL

Swimmera movement

OVl-H STRENGTH

FIGURE 1. View of the swimmers situations during theexercises in overstrength and in overspeed.

ProceduresSwimmers were randomly divided into 3 groups: (a) over-strength (resisted-sprint) group fOSt; n = 15; age 16.5 ±2 years, height 170 ± 7 cm, weight 58 ± 9.5 kg), (b) over-speed (assisted-sprint) group (OSp; n ^ 11; age 18 ± 3years, height 176 ± 7 cm, weight 67 ± 10.5 kg), and (c)control group (C; n = 11; age 17 ± 3 years, height 168 ±9 cm, weight 62 ±11 kg).

The training program lasted 3 weeks during the Feb-ruary month, at the beginning of the second macrocycleof the season. Average training volume and intensitywere the same for all swimmers throughout the studyprotocol. A total of 10 sessions per week, from Monday toFriday (2 sessions per day, 1 in the morning and 1 in theevening, 1.5 hours per session) were performed duringthe training period. During the 3 weeks, the training vol-ume was 45 ± 5 km per week. The training program in-cluded a dominant aerobic work in front crawl during themorning training sessions and a dominant technical workin medley during the evening training sessions, at a mod-erate intensity.

Swimmers in the OSt group swam 6 all-out 30-secondfront crawl sprints with a 30-second recovery period be-tween each sprint (total duration 6 minutes). The swim-mers were tethered to the starting platform. They wore abelt around the pelvis that was attached to one end ofa 5-m-long elastic surgical tube (Paul Factory, Saint-Etienne, France; inner and outer diameters 8 and 12 cm,respectively); the other end of the tube was attached tothe starting platform (Figure 1). The elastic tube imposeda length (Y: in meters)~strength {X; in Newtonsf relation-ship on the swimmers expressed by Y 4.5008 X -12.95.The elastic stretched over an average distance of 15 m.During recovery, the swimmers held onto the swimminglane to keep the tuhe taut.

Swimmers in the OSp group swam 12 25-m freestylefront crawl sprints. These swimmers were also attachedto an elastic surgical tuhe under the same conditions asfor OSt except that the tube was 8 m long and attachedto the point of arrival. The point of departure was set atthe 25-m line so that the elastic tube pulled the swimmertoward the point of arrival with an initial force of 60 N.As the swimmer advanced, an assistant maintained the

elastic as taut as possible to maintain the same forcethroughout the sprint. The swimmers were asked to fol-low the speed given by the elastic by having a high strokerate and trying to not decrease their distance per stroke.Between each sprint, the swimmer got out of the pool,walked back to the point of departure, and jumped intothe water. This walk back was considered as a recoveryperiod so the total duration was about the same as for theOSt group (about 6 minutes).

The control group (C) swam 6 50-m all-out front crawlsprints, without an elastic tube, with a 30-second recov-ery period between sprints, for a total duration of about6 minutes.

The training sets described above were the only dif-ferences in training hetween the 3 groups. These setswere performed 3 times per week, in the first part of thetraining sessions, after a 1,500-m standardized warm up,on Monday, Wednesday, and Friday evening of each week(Tahle 1). No dry-land training program was performedduring the 3 weeks. Swimmers were asked to refrain fromany other sport activity.

Muscle Strength MeasurementsThe flexion-extension peak torque of the 2 forearms, ex-pressed in newton-meters (N-m), was measured with anisokinetic dynamometer (Cybex; Medimex Factory, Tas-sin la Demi Lune, France) before the training program(week 0) and after 2 (week 2) and 3 (week 3) weeks oftraining. Forearm peak torque was retained for study be-cause measurement is easier than with the whole arm,and, as demonstrated by Shleihauf et al. (22), forearmforces account for a large part of total arm propulsion.Before the measurements, a 5-minute standardized warmup and familiarization period was performed with the ap-paratus at several submaximal intensities (60", 180", and0°-s '). These different angular velocities have been cho-sen because they seemed to be most representative of aswimmer's movement speed, as reported by Pichon et al.(19). The measurements took place at the end of eachweek, on a Saturday afternoon (24 hours after the lasttraining session in assisted sprint). Swimmers sat withthe torso strapped at the shoulders and pelvis. The armwas maintained parallel to the Cybex's arm lever. Thespindle of the motor was positioned in line with the centerof rotation of the elbow joint. The measurements weremade under concentric conditions at 60° and 180°-s ' andunder isometric conditions at O' -s '. The subjects wereasked to perform 2 maximal efforts with 3 repetitions al-ternating flexion and extension at 60° s ' and at 180°-s '.The best performance was retained. A 30-second rest pe-riod separated each effort, and a 2-minute rest period sep-arated each velocity. In isometric condition, the subjectswere asked to perform 2 maximal efforts in flexion andin extension. The efforts lasted 5 seconds with a 2-minuterest period between repetitions; the elbow angle betweenthe arm and forearm was set at 90''. Intraclass correla-tions (ICC R) of the physical strength measurements, as-sessed in 18 swimmers using the coefficient of variationof the difference between 2 measurements, were 4.2%.

Swimming Performances and Technical ParametersSwimming performances were measured at weeks 0, 2,and 3, the same weekday at the same time of the day,during a 100-m front crawl competition in a 25-m poolafter a 30-minute warm up. The 100-m event was chosen

ASSISTED AND RESISTED SPRINT TRAINING IN SWIMMINC; 549

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for study because it is the most common sprint distancein competition, midway between the 50-m short sprintand the 200-m long sprint. In addition, it is generally themost popular distance for swimmers. All the races werevideo recorded. The stroke rate, expressed as the numberof strokes per minute, was measured twice every 50 m,once every 25 m, using a stroke base 3 stopwatch (Seiko;Seiko Factory, Besangon, France). The average value wasretained. The distance per stroke was calculated by di-viding the mean velocity of each 50 m by the mean strokerate.

Statistical Analyses

Results are presented with their mean and SD values.Analyses of variance were used to compare the main char-acteristics, performances, muscular strength, stroke rate,and distance per stroke ofthe 3 groups. A Tukey-Kramertest was used as a post hoc test. Correlation coefficientswere calculated between the performance and the differ-ent measured parameters. For the whole group stepwiseregressions were calculated between the 100-m frontcrawl velocity (independent variable) and the other vari-ables {dependent variahle) using the StatView 512+ pro-gram (SAS Institute Inc., Cary, NC). In all ofthe statis-tical analyses, significance was accepted at p < 0.05.

RESULTS

Before training, there was no significant difference be-tween stroke rates and distances per stroke for the 3groups (Table 2). However, between OSt and OSp groups,there was a significant difference in body weight, perfor-mance ip < 0.05, Table 2), and elbow extensors isometric(O^-s') and isokinetic (60°-s ') strength ip < 0.05, Figure2). For the entire study population, 100-m performancewas related to the strength of the elhow flexors and ex-tensors under isometric conditions (r = 0.57; 0.54; p <0.05) and concentric conditions at 60° s ' {r - 0.67; 0.66;p < 0.05) and 180°-s ' (r = 0.64; 0.66; p < 0.05). It wasalso related to stroke rate (r = 0.45; p < 0.05) and strokelength ir = 0.48; p < 0.05). These relationships were alsofound in each of the 3 study groups.

Effect of Training on Muscle Strength

The effect of training on muscle strength is presented inTahle 3 and Figure 2. For OSt, muscle strength was sig-nificantly (p < 0.05) increased in 3 conditions: the iso-metric condition for the elbow extensors and the 2 con-centric conditions for the fiexors ip < 0.05). For OSp,muscle strength was significantly (p < 0.05) increasedonly in 1 condition: the concentric condition at lSO -s for the fiexors. For the control group (C), muscle strengthwas significantly ip < 0.05) increased in the isometriccondition (O^'-s'), for the fiexors.

Data from week 2 (not presented) did not further mod-ify the effect of training on muscle strength. Data pre-sented in Table 4 were obtained after 3 weeks of training.

Effect of Training on Swimming Performance

For OSt, the gain in swimming performance was signifi-cant (p < 0.05) over the whole training period (Table 2),while for OSp it was significant (p < 0.05) only hetweenthe second and third weeks. For the 2 groups (OSt andOSp), the gain was significant during the second 50 m ofthe 100-m swim (p < 0.05) but not during the first 50 m.

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No significant difference was observed for the controlgroup.

Effect of Training on Technical ParametersFor OSt and OSp, stroke rate was significantly (p < 0.05)increased over the whole training period (Table 2). Thegain was significant only during the second 50 m of the100-m swim (p < 0.05). No significant difference was ob-served for the control group. For OSt, the distance perstroke was maintained during the second 50 m (Table 2),while for OSp, it decreased significantly (p < 0.05). Nosignificant difference was observed for the control group,Considering all swimmers together, stroke rate varia-tions, expressed in percentage of baseline, correlated withperformance variations, expressed in percentage of base-line performance (r = 0.68; p < 0.05; Figure 3). Theserelationships were also found in each of the 3 studygroups; for the OSt (r = 0.65; p < 0.05), for the OSp (r =0.73; p < 0.05), and for the C (r = 0.59; p < 0.05).

Training Effect: Comparison between Strength andTechnical ParametersConsidering all swimmers, stepwise regression analysisbetween swimming performance and physical strengthand technical parameters revealed that stroke rate dur-ing the second 50 m (SR 2nd 50) was the only factor ex-hibiting significant correlation (r ^ 0.68; p < 0.05).

For the OSt group, SR 2nd 50 was the most significantfactor, while physical strength ofthe elbow flexors in con-centric conditions at 60" s ' (ISA-60) was the second mostimportant factor. The effects of these 2 factors was addi-tive, the effect of iSA-60 significantly increasing the co-efficient of correlation between performance and SR 2nd50 from 0.77 to 0.84 (p < 0.05), according to the followingequation:

^Performance - (0.263 x SR 2nd 50)

+ (-0.91 X ISA-60) + 1.321

For the OSp group, stroke rate during the first 50 m(SR 1st 50) was the most significant factor, while strokelength during the first 50 m (SL 1st 50) was the secondmost important factor. The effects of these 2 factors wereagain additive, significantly increasing the correlation co-efficient from 0.62 to 0.82 (p < 0.05), according to thefollowing equation:

APerformance = (0.13 X SR 1st 50)

+ (0.328 X SL 1st 50) + 0.827

For group C, SR 2nd 50 was the most significant fac-tor, while physical strength of the elbow flexors in con-centric conditions at 180°-s ' (lSA-180) was the secondmost important factor. The coefficient of correlation in-creased significantly from 0.73 to 0.88 (p < 0.05), accord-ing to the following equation:

APerformance - (0.149 X SR 2nd 50)

+ (-0.63 X ISA-180) + 0.824

Gender EffectIn each group, there was a significant difference betweenmen and women in performance before training (p <0.05). In the OSt group, there were significant differencesin training effects between men and women on perfor-mances and stroke rate (p < 0.05). In this group, only

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FIGURE 2. Relationship between the peak torque and the angular velocity before and after training for the different groups forthe elbow extensors and flexors. *Significance ip < 0.05). Values are mean and SEE. For the extensors, overstrength group = A,overspeed group = B, control group = C. For the flexors, overstrengtb group ^ D, overspeed group ^ E, control group = F.

TABLEvelocities

3. Comparison of the variations of the physical strength, expressed in percentage of the initial values.*

Flexors

Isometric

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' 180"s

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14.2 (27.2)

3.7 (10.7)

141 (15.3)

21.It (23.1)

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15.3 (18.3)

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10.5 (11.9)

1.5 (10.2)* Values are mean (± SD); n, number of subjects; significant at p < 0.05.t Significantly different from pretraining values.

vyomen increased significantly in performances and instroke rate {p < 0.05). There were no significant differ-ences in training effects betvifeen men and women in theother groups.

D I S C U S S I O N

The main findings of the present study were (a) the re-sisted-sprint training program was more efficient thanthe assisted-sprint program in increasing musclestrength and lOG-m performance and (b) physicalstrength and technical parameters were good predictorsof 100-m performance.

The resisted-sprint program was the most efficienttraining method to increase muscle strength after 3weeks of training when compared with the assisted-sprintprogram and standard training (control group). The sig-nificant increase in the strength of the elbow extensorsmeasured in isometric conditions was 32% in the OSt

group vs. 5% and 15% for the OSp and control groups,respectively (Table 3). These results are close to the 24%gain for 3 weeks of training reported by Pichon et al. (19)and to the 20-40% gain reported by Strass et al. (26). TheOSt group also exhibited a 2% increase in 100-m perfor-mance compared witb 0.8% for the OSp group and -0.3%for the C group. These data do not corroborate those re-ported by Delecluse et al. (8, 9) who worked with runnersand found that the assisted-sprint training method wasthe most efficient. In their studies, the gain in runningperformance was related to a gain in departure reactiontime but not in muscular strength. Departure reactiontime was not measured in the present study, but for a100-m swim (taking about 1 minute) departure time isrelatively less important tban tbat for a 10-12-second100-m run, and may explain the observed differences. Inthe OSt group, the fact that females significantly in-creased their performance, and not males, was probably

552 GiROLD, CAI.MELS, M A U R I N ET AL.

FIGURE 3. Relationship between the variations ofperibrniance and the variation of stroke rate during the second50 m of the 100-m swim for the whole study population.

due to differences in their levels at the heginning of thetraining program. As males were significantly faster thanfemales in the 100-m distance hefore training, it washarder for them to significantly enhance their speed thanit was for females who had a lower level.

Muscle strength and technical parameters were goodpredictors of 100-m swimming performance. Strength ofelbow flexors and extensors was correlated with 100-mperformance measured in isometric and concentric con-ditions. These data are in agreement with previous stud-ies reported by Hawley and Williams (11, 12) and Mi-yashita and Kanshisa {17} showing a strong relationshipbetween the power of the upper limbs and 100-m swim-ming performance. In the present study, strength gainwas greater in isometric conditions. Under these condi-tions, the arm-forearm angle is 90°. This angle corre-sponds to the specific mid-stroke swimming position, be-tween tbe pulling and pushing phases (16). In this posi-tion, the arm is deep in the water and requires greaterpower to overcome the resistance of the water. Thus, again in muscle strength in this position may be more ef-ficient in terms of performance gain. The present dataalso confirmed work by Costill (7) and Stewart and Hop-kins (25) emphasizing the importance ofthe specificity oftraining methods in improving swimming efficiency.

The gain in muscular strength was greatest for theelbow extensors, basically the triceps brachii. Birrer (3)also painted out the importance ofthe triceps in the push-ing phase for all strokes, and Rouard et al. (20) andSchleihauf et al. (22, 23) found that peak force occurs atthe end ofthe aquatic phase ofthe stroke during forearmextension on the arm.

Two technical parameters, stroke rate and strokelength, were related to swimming perfonnance, confirm-ing reports by Arellano et al. (1), Hohmann et a!. (13),Keskinen et al. {15), and Sidney et al. (24) indicating tbattechnical parameters are directly related to sprint perfor-mance. Considering the entire study population, musclestrength was more important than technical parametersas demonstrated by the higher coefficient of correlationbetween muscle .strength and performance ir = 0.67) thanbetween technical parameters and performance (r ^0.48). However, stepwise regression analysis indicatedthat tbe most important factor directly related to gain inperformance was stroke rate and not muscle strength(Figure 3). These results are probably related to the du-

ration of the training program, which may have been longenough for nervous system adaptation to induce in-creased speed of movement hut not long enough for thedevelopment of sufficient physical strength to have a de-termining effect on gain in performance.

Swimmers in the OSt group increased their strokerate solely in the second part of the 100-m swim whilethe swimmers in the OSp group increased their strokerate in both parts. But gain in performance was signifi-cant only for tbe OSt swimmers because they maintainedtheir distance per stroke while the OSp swimmers did not(stroke length decreased in the OSp group). This findingis in agreement with the previous study of Maglischo etal. (16) indicating that assisted-sprint swimming exercis-es cause a greater increase in stroke rate than resisted-sprint swimming exercises. Maglischo et al. (16) observedthat assisted-sprint swimming exercises also cause agreater decrease in stroke length than tethered-sprintswimming exercises. Delecluse et al. (8) suggested that ahigh-resistance and high-velocity sprint training programenhances power and movement speed due to adaptivechanges in the nervous system. Nevertheless, adaptationsresulting from the 2 training methods would be different.Delecluse et al. (8) suggested that a high-resistance sprinttraining program first develops motor unit recruitmentcombined with a gain in movement velocity, but that gainin velocity is limited by the time maximal motor unit re-cruitment takes to generate maximal strength. A high-velocity sprint training program first develops movementspeed (despite the movement length), combined with again of power at high velocity. This could explain why theOSt (resisted-sprint) group exhibited increased physicalstrength in isometric conditions, while the OSp (assisted-sprint) group developed increased physical strength inconcentric conditions at 18O''-s ' (Table 4), as well as thedifferences in stroke rate and stroke length gains andvariations between the 2 groups.

PRACTICAL APPLICATIONS

The present study indicated that, in a sprint training pro-gram, resisted-sprint training is more efficient than as-sisted-sprint training to achieve increased performance in100-m front crawl swimming. The observed gain was ex-plained by an increase in stroke rate. Physical strengthwas modified by both training methods at different an-gular velocities. Further investigations are required todetermine the underlying muscular or neurologicalchanges involved in these adaptations.

These training methods can be used all along tbe sea-son. In a period of high training volume, resisted-sprintcan develop strength endurance in the water while as-sisted-sprint may increase the hydrodynamic position andthe stroke rate. In this period it must be realized withlong sets at a moderate intensity with a short recoverytime. In a period of competition, resisted-sprint can beused to increase strength and power, and assisted-sprintto increase stroke rate and strength at a high velocity. Inthis period, it must be realized with short sets at a max-imal intensity with a long recovery time.

The corresponding time ofthe swimming velocity gainover 100 m after the 3-week training period was: 1.9 ±0.9 seconds in OSt, 1.1 ± 0.8 seconds in OSp and 0.5 ±0.9 seconds in C. The authors had realized another study(unpublished data) over a period of 12 weeks in resisted-and assisted-sprint. After this period, swimmers in-

ASSISTED AND RESISTED SPRINT TRAINING IN SWIMMINCI 553

TABLE 4. Comparison ofthe 100-m performances divided in two 50 m.*

Group

Overstrength group{n = 15)

Overspeed group(n = 11)

Control group(n - 11)

1st 50 m(s)

32.28§(2.34)29.86|{(2.72)32.64(3.02)

WeekO

2nd 50 m(s)

35.22§(2.18)32.5911(2.66)35.51(3.27)

100 m(s)

67.43§(4.4)62.4611(5.32)68.15(6.18)

1st 50 m(s)

32.07t§(1,92)29.9311(2.68)32.49(2.79)

Week 2

2nd 50 m(s)

35.20t§(2.10)32.75t||(2.78)35.89(3.34)

100 m(s)

67.14t§(3.95)62.67tll(5.45)68.39(6.09)

1st 50 m(s)

31.75§(2.05)29.6911(2.45)32.85(2.77)

Week 3

2nd 50 m(s)

34.30$§(2.01)

32.251:11(2.45)35.50(3.17)

100 m(s)

66.05|ij(4)61.911(4.85)68.35(5.91)

*• Values a re mean (± SD); n, n u m h e r of subjects; significant a t p < 0.05.1" Significant difference hetween 2nd and 3rd 100 m.:1: Significant difference between I s t and 3rd 100 m.§ Significant difference hetween overs t rength and overspeed.II Significant difference be tween overspeed a n d control,

creased significantly their performances by 3.5% over 50m vs. 2% in the present study over 100 m and after only3 weeks of training. Thus, it can be supposed that thelonger the training period, the stronger the effect on per-formances. However, performance improvements are al-ways greater when new training methods are proposed,as their efficiencies decrease with time.

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Acknowledgments

The authors thank the swimmers for their voluntaryparticipation in the training program. Prof, Jacques Duchateaufor making valuahle suggestions, Gilhert Lomhard for histechnical assistance, and Gerald Pope for reviewing the Englishlanguage of the manuscript-

Address correspondence to Dr. Sebastien Girold,xgirold@free.fr.