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Muscle Swelling of the Triceps Surae in Response to Straight-Leg and Bent-Leg

Calf Raise Exercises in Young Women

Article in The Journal of Strength and Conditioning Research · July 2023

DOI: 10.1519/JSC.0000000000004491

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1 Muscle swelling of the triceps surae in response to straight-leg and bent-leg calf raise

2 exercises in young women

3 Short title: Different calf raise exercises and muscle swelling

4 Authors: Witalo Kassiano1, Bruna Costa1, Gabriel Kunevaliki1, Danrlei Soares1, Natã Stavinski1,

5 Jarlisson Francsuel1, Marcelo A. S. Carneiro1, Ian Tricoli1, João Pedro Nunes1,3, Alex S. Ribeiro2,

6 Edilson S. Cyrino1

7 Affiliations: 1Metabolism, Nutrition and Exercise Laboratory, Physical Education and Sport

8 Center, State University of Londrina, Londrina, PR, Brazil; 2University Pitágoras UNOPAR. Lon-

9 drina, PR, Brazil; 3Centre for Human Performance, School of Medical and Health Sciences, Edith

10 Cowan University, Perth, Australia.

11

12 *Correspondence author: Witalo Kassiano. e-mail: acc.witalo@gmail.com | ORCID number:

13 0000-0002-0868-8634. Metabolism, Nutrition, and Exercise Laboratory. Physical Education and

14 Sport Center, State University of Londrina, Rodovia Celso Garcia, km 380, 86057-970, Londrina,

15 PR, Brazil.

16

17 Affiliation where the research was conducted: Physical Education and Sport Center. State

18 University of Londrina, Brazil.

19 ABSTRACT

20 Triceps surae muscle swelling in response to different resistance exercises remain to be deter-

21 mined. The present study compared the effects of straight-leg (STRA-leg) calf raise versus bent-

22 leg (BENT-leg) calf raise exercises on triceps surae muscle swelling. Seventeen young women

23 (23.7 ± 4.0 yrs; 67.4 ± 16.0 kg; 163.5 ± 7.2 cm) performed two resistance training sessions; in one

24 they performed the STRA-leg calf raise and in another, they performed the BENT-leg calf raise.

25 A randomized, crossover, and counterbalanced design was adopted for this investigation. The sub-

26 jects performed 4 sets of 20 repetitions maximum until concentric muscular failure. The muscle

27 thickness of gastrocnemius medial (GM), gastrocnemius lateral (GL), and soleus (SOL) were taken

28 via B-mode ultrasound pre and immediately post the calf raise exercises. STRA-leg calf raise elic-

29 ited greater increases in muscle thickness of GM (+8.8% vs. -0.9%, P < 0.001) and GL (+14.5%

30 vs. +7.0%, P < 0.001) than BENT-leg calf raise. Conversely, BENT-leg calf raise elicited greater

31 increases in SOL muscle thickness than STRA-leg calf raise (+15.4% vs. +7.7%, P < 0.001). From

32 a practical perspective, the STRA-leg calf raise should be preferred if the aim is stimulating the

33 three muscles that comprise the triceps surae, whereas the BENT-leg calf raise should be priori-

34 tized if the target muscle is the SOL.

35 Keywords: resistance exercise, muscle thickness, ultrasound, exercise selection, ankle flexors,

36 active insufficiency.
37 INTRODUCTION

38 Resistance exercises induce inhomogeneous stimulus along the length of a muscle and be-

39 tween muscles of a muscle group (16, 19, 30). For instance, immediately after performing the

40 barbell back squat, there is an increase in the cross-sectional area of the vastii and adductor but not

41 in the rectus femoris (19). This transient augment in muscle size has been termed muscle swelling

42 and appears to occur due to a fluid shift to exercised muscles due to metabolic demand in response

43 to repeated muscle contractions (5, 21). Based on this, muscle swelling has been considered a valid

44 proxy to identify which muscles are most stimulated in response to resistance exercise (5). Inter-

45 estingly, there appears to be a relationship between muscle swelling measures and muscle hyper-

46 trophy (12). Ultrasound imaging has been used to detect changes in muscle size after exercise (12,

47 26, 27). However, most studies investigating resistance exercise-induced muscle swelling have

48 focused on the quadriceps (12, 26, 27). Therefore, findings characterizing swelling effects of dif-

49 ferent resistance exercises for other muscles such as triceps surae are limited or non-existent.

50 The triceps surae is a muscle group that arouses the interest of strength and conditioning

51 scientists and practitioners, in part, because it is deemed as difficult to respond to hypertrophy-

52 oriented resistance training programs (3, 17). This large muscle group consists of the gastrocnem-

53 ius medial (GM), gastrocnemius lateral (GL), and soleus (SOL) (2, 14). The SOL is a single-joint

54 plantar flexor muscle that crosses the ankle exclusively. In contrast, the gastrocnemius heads are

55 two-joint muscles that cross the knee and ankle joints (14). Based on this anatomical feature, gas-

56 trocnemius heads may experience active insufficiency at greater knee flexion angles (14). There-

57 fore, reducing its contribution, for example, during a calf raise exercise performed with the bent-

58 leg. Theoretically, this would result in a greater contribution of the SOL in a bent-leg (BENT-leg)

59 calf raise (22). On the other hand, the calf raise performed with the straight-leg (STRA-leg) could
60 favor a greater contribution—therefore, superior stimulus—for the GM and GL, as it is in a favor-

61 able force-tension relationship, consequently, more favorable force-generating capacity (14).

62 Despite the plausibility of the arguments presented above, the evidence is inconclusive.

63 For instance, a previous study (24) reported greater SOL activation (measured via surface electro-

64 myography [sEMG]) with the knees bent at 90º than at 0º (i.e., fully extended). On the other hand,

65 another investigation detected similar sEMG amplitude in the SOL with knees at 90º and 0º (7).

66 Moreover, sEMG amplitude in the GL was not impaired in function of knee angle (24). In contrast,

67 another report (7) detected lower sEMG amplitude in the GL at 90º of knee flexion. These diver-

68 gences may occur due to inconsistencies related to the measurement method (i.e., sEMG), partic-

69 ularly when comparing different muscle lengths (28, 29). In this context, muscle swelling may be

70 an alternative method to be used to represent muscle stimulation. Thus, this study compared swell-

71 ing of the triceps surae muscles immediately after STRA-leg and BENT-leg calf raise exercises.

72 We hypothesized that the gastrocnemius heads muscle swelling would be greater in the STRA-leg

73 calf raise, whereas SOL muscle swelling would be greater in the BENT-leg calf raise exercise.

74

75 METHODS

76 Experimental approach to the problem

77 A randomized, crossover, and counterbalanced design was adopted for this investigation. The total

78 duration of the study was 4 weeks. Weeks 1 and 2 were used for anthropometry measurements and

79 repetitions maximum (RM) testing on straight-leg and bent-leg calf raise exercises. Weeks 3 and

80 4 were used for the application of the experimental training sessions. Subjects performed the train-

81 ing sessions with straight-leg (STRA-leg) or bent-leg calf raise (BENT-leg) exercises following a

82 randomized, counterbalanced order; 9 performed the STRA-leg calf raise in the first training
83 session and 8 performed the BENT-leg calf raise in the first training session. In the 2 experimental

84 training sessions, we measured the muscle thickness of GM, GL, and SOL before and immediately

85 after the last set via ultrasonography. We recorded the load used and the number of repetitions and

86 repetitions in reserve following each set. A 1-week washout period was given between experi-

87 mental training sessions. All subjects performed their RM tests and training sessions at the same

88 time of the day (2 p.m. to 6 p.m.) to minimize the possible effects of the circadian cycle on strength

89 performance. Subjects were instructed to maintain their nutritional habits and hydration and did

90 not engage in resistance exercise during their participation in the present study.

91 Subjects

92 Seventeen recreationally active young women (23.7 ± 4.0 yrs, 67.4 ± 16.0 kg, 163.5 ± 7.2 cm)

93 participated in the present study. The sample size was determined a priori using G*Power software

94 (version 3.1.9.6) with the significance level set at 0.05 and power of 0.90 to detect a moderate

95 effect size (F = 0.29) (26). The analysis indicated at least 16 subjects were needed to achieve

96 adequate statistical power. Volunteers were recruited through dissemination on social media and

97 folders distributed in points of greater circulation within the campus of the local University. All

98 subjects had experience with resistance training but had not trained for 2 months before the start

99 of the present study. Notably, the participants had familiarization with the straight-leg and bent-

100 leg calf raise exercises. Before the inclusion, they completed a detailed health history questionnaire

101 and were included in the study if they had no signs or symptoms of disease, were not using medi-

102 cations or supplements, and had no orthopedic injuries. Written informed consent was obtained

103 from the subjects after a detailed description of all procedures. This investigation was conducted

104 according to the Declaration of Helsinki and was approved by the local University Ethics Com-

105 mittee (protocol number: 3.930.966).

106 Procedures

107 Triceps Surae Muscle Thickness Measurement. Muscle thickness measurements of the GM, GL,

108 and SOL were assessed using a B-mode ultrasound imaging device (FIGLABS, model FP-102,

109 SAEVO, São Paulo, SP, Brazil), with a 53 mm, 7.5-MHz linear probe model L741. The subject

110 was placed in a prone lying position on a stretcher and rested for 10 min before the beginning of

111 the assessment. Images were obtained on the posterior surface of the right leg with the probe per-

112 pendicular to the tissue. Image acquisitions of the GM were taken with the probe positioned in the

113 thickest and more prominent site of the leg from a posteroanterior view (17). Measurements of the

114 GL were taken at the proximal third between the femur lateral epicondyle and the fibula lateral

115 malleolus (Figure 1A). Image acquisitions of the SOL were taken at the middle distance between

116 the femur and the lateral malleolus of the fibula (Figure 1A). Reference lines were drawn on the

117 subject’s skin with a dermatographic pen on the sites at which images were obtained. The lines

118 were reinforced and maintained throughout the study to ensure that the measurement was taken at

119 the same site pre- and post-exercise and in the 2 experimental training sessions. A generous quan-

120 tity of water-soluble transmission gel was applied over the muscle being assessed, without com-

121 pressing the skin. Two assessors participated in measurement procedures. The first handled the

122 probe, and the second was responsible for freezing the images. When the image quality was

123 deemed to be satisfactory, the second assessor, who was blinded to condition allocation, obtained

124 muscle thickness dimensions. Muscle thickness of GM and GL was defined as the distance from

125 the superficial to deep aponeuroses that borders the SOL (Figure 1B). The SOL was defined as the

126 distance from the upper and lower aponeuroses separating the muscle (Figure 1B). The muscle

127 thickness dimensions were obtained on the ultrasound machine’s calculation package through a

128 line drawn at right angles in the center of each ultrasound image. Eleven subjects were randomly
129 chosen to be evaluated on two days separated by 72 h to determine the reliability of the muscle

130 thickness measurements. The intraclass correlation coefficients for the GM, GL and SOL were

131 0.978, 0.989, and 0.987; the coefficients of variation were 2.1%, 2.5%, and 1.3%; the standard

132 errors of measurement were 0.086 cm, 0.074 cm, and 0.077 cm, respectively. Measurement of

133 muscle thickness after the training session was started less than 1 minute after the end of the last

134 training set. This was possible by taking the ultrasound device and a stretcher to the gym room

135 where the training sessions (described below) were performed.

136

137 *** PLEASE INSERT FIGURE 1 NEAR HERE ***

138

139 Repetitions Maximum Test. The determination of the weight to be used in STRA-leg and BENT-

140 leg calf raise exercises was performed through RM testing. Initially, the subjects performed a

141 warm-up in the specific exercise with half the weight that would be used on the first set (selected

142 based on researcher and subject experience). After 1 minute of rest, the first set was started. The

143 subjects were asked to perform 20 repetitions in both exercises. This number of repetitions was

144 chosen based on its common inclusion in calf hypertrophy-oriented training programs (13, 17). If

145 the subjects were able to perform 20 repetitions without reaching the task failure, the weight was

146 increased by 2–5% for the next set. On the contrary, if the subject could not perform 20 repetitions,

147 the weight was decreased by the same magnitude. Each subject performed three sets per testing

148 session with a 3-minute rest interval. An interval of 20 min was given between the RM testing in

149 BENT-leg and STRA-leg calf raise. The RM testing was performed on two days, separated by 48

150 h. The weight with which the subject could not exceed 20 repetitions was used in the experimental

151 training sessions. The intraclass correlation coefficients for the RM testing in STRA-leg and
152 BENT-leg calf raise were 0.988 and 0.980; the coefficients of variation were 1.7% and 2.7%; the

153 standard errors of measurement were 2.41 kg and 1.52 kg, respectively.

154 Experimental Training Sessions. Two experimental training sessions were performed; in one the

155 subjects performed the STRA-leg calf raise exercise and in another, they performed the BENT-leg

156 calf raise exercise. The 2 experimental conditions were carried out in a randomized and counter-

157 balanced order with a 1-week washout between each one. STRA-leg calf raise exercise was per-

158 formed with the knees at 0º (i.e., fully extended), in a pin-loaded horizontal leg-press machine

159 (Ipiranga®, Presidente Prudente, Brazil) (Figure 2A). BENT-leg calf raise exercise was performed

160 with the knees at 90º, in a specific device (Ipiranga®, Presidente Prudente, SP, Brazil) with weight

161 plates (Figure 2B). The subjects performed a warm-up set of 20 repetitions with 50% 20RM, and

162 after 1-min of rest, the subjects performed 4 sets with 20RM load. Subjects were verbally encour-

163 aged to achieve task failure in each set—i.e., the point at which, despite trying to do so, they cannot

164 complete the concentric muscle action of their current repetition (25). A 2-min of rest interval was

165 given between sets. Exercises were performed in the full range of motion, in a tempo of 1/0/2/1

166 (concentric, concentric-eccentric transition, eccentric, and eccentric-concentric phases, respec-

167 tively). The foot was positioned on the platform supported by metatarsals. The number of repeti-

168 tions in each set was recorded. After each set, the number of repetitions in reserve (RIR) was

169 quantified. After each exercise set, subjects answered the following question: “How many addi-

170 tional repetitions could you have performed?” (11). An estimated repetition to failure score of “0”

171 indicated that the subject estimated that no additional repetitions could be completed (0 RIR; task

172 failure reached), “1” = 1 RIR; “2” = 2 RIR, and so forth.

173

174 *** PLEASE INSERT FIGURE 2 NEAR HERE ***

175

176 Statistical analysis

177 The normality of the data was tested using the Shapiro-Wilk. The number of repetitions and RIR

178 were compared using a repeated-measures analysis of variance (ANOVA) with condition (STRA-

179 leg calf raise vs. BENT-leg calf raise) × time (1st, 2nd, 3rd, and 4th sets) as fixed factors. When

180 sphericity was violated, the Greenhouse-Geisser correction factor was applied. The absolute load

181 used in the exercises was compared using an independent t-test. Muscle thickness of the GM, GL,

182 and SOL were compared using a repeated-measures ANOVA with the condition (STRA-leg calf

183 raise vs. BENT-leg calf raise) × time (pre- vs. post-session) as fixed factors. When the F was

184 significant, a Bonferroni post-hoc test was used to identify possible statistical differences. The

185 effect size (ES) was calculated as post-mean minus pre-values mean, divided by pooled pre-values

186 standard deviation (6). The data were presented in mean, standard deviation, and 95% confidence

187 interval. The accepted level of significance was P < 0.05. The data were analyzed using the JASP

188 software (version 0.15, Amsterdam, NL).

189

190 RESULTS

191 The absolute load was significantly greater in the STRA-leg than in the BENT-leg calf

192 raise (51.0 ± 12.0 kg vs. 23.5 ± 5.0 kg; t = 11.317, P < 0.001). The number of repetitions and RIR

193 are presented in Table 1. There was a significant time effect (F = 116.800, P < 0.001), but no

194 condition (F = 0.936, P = 0.348) or time × condition interaction (F = 0.439, P = 0.726) for the

195 number of repetitions. The number of repetitions was lower in the 3rd and 4th sets than in the 1st

196 and 2nd sets in both training sessions. There was a significant main effect of time (F = 4.745, P =

197 0.006), but no condition (F = 0.314, P = 0.583) or time × condition interaction (F = 0.584, P =
198 0.628) for the number of RIR. The number of RIR was lower in the 2nd, 3rd, and 4th sets than in the

199 1st set in both training sessions.

200

201 *** PLEASE INSERT TABLE 1 NEAR HERE ***

202

203 Table 2 displays the pre- and post-session values of the GM, GL, and SOL muscle thick-

204 ness. There was no significant difference in the pre-sessions muscle thickness between the exper-

205 imental conditions (all P > 0.05). Figure 3 shows the individual changes in muscle thickness of the

206 GM, GL, and SOL.

207

208 *** PLEASE INSERT TABLE 2 NEAR HERE ***

209 *** PLEASE INSERT FIGURE 3 NEAR HERE ***

210

211 For the GM, there was a significant main effect of time (F = 27.898, P < 0.001), condition

212 (F = 45.629, P < 0.001), and time × condition interaction (F = 61.795, P < 0.001). There was

213 increase in GM muscle thickness pre-to-post STRA-leg calf raise (∆ = +8.8%, P < 0.001), but not

214 pre-to-post BENT-leg calf raise (∆ = -0.9%, P = 0.999). GM muscle thickness was significantly

215 greater at post STRA-leg than post BENT-leg calf raise (Meandiff = 0.21 cm, 95% CI: 0.15, 0.27;

216 P < 0.001).

217 For GL, there was a significant main effect of time (F = 42.000, P < 0.001), condition (F

218 = 11.755, P = 0.003), and time × condition interaction (F = 12.919, P = 0.002). There was increase

219 in GL muscle thickness pre-to-post STRA-leg calf raise (∆ = +14.5%, P < 0.001), and pre-to-post
220 BENT-leg calf raise (∆ = +7.0%, P = 0.013). GL muscle thickness was significantly greater at post

221 STRA-leg than post BENT-leg calf raise (Meandiff = 0.13 cm, 95% CI: 0.05, 0.21, P < 0.001).

222 For SOL, there was a significant main effect of time (F = 273.432, P < 0.001), condition

223 (F = 5.559, P = 0.031), and time × condition interaction (F = 31.348, P < 0.001). There was an

224 increase in SOL muscle thickness pre-to-post STRA-leg calf raise (∆ = +7.7%, P < 0.001), and

225 pre-to-post BENT-leg calf raise (∆ = +15.4%, P < 0.001). SOL muscle thickness was significantly

226 greater at post BENT-leg than post STRA-leg calf raise (Meandiff = 0.14 cm, 95% CI: 0.06, 0.22,

227 P < 0.001).

228

229 DISCUSSION

230 The purpose of the present study was to compare the acute effects of performing STRA-

231 leg and BENT-leg calf raise on muscle thickness of the triceps surae muscles. The main findings

232 of this study were that exercise selection influences the muscles that experience muscle swelling

233 and the magnitude of this transient increase in muscle size. Specifically, STRA-leg calf raise elic-

234 ited muscle swelling in the three triceps surae muscles, whereas BENT-leg calf raise induced mus-

235 cle swelling in the GL and SOL, but not in GM. Notably, the magnitude of muscle swelling on

236 gastrocnemius muscles was greater after the STRA-leg than the BENT-leg calf raise. In contrast,

237 the magnitude of muscle swelling on the SOL was greater after the BENT-leg than after the STRA-

238 leg calf raise. It is important to highlight that such effects occurred with the calf raise exercises

239 being performed with the same relative load (i.e., 15-20RM) and similar proximity to task failure

240 (i.e., 0 RIR). Our initial hypothesis that STRA-leg calf raise would be induced superior gas-

241 trocnemius muscle swelling but inferior SOL muscle swelling than BENT-leg calf raise was con-

242 firmed.
243 In the present study, we observed that the magnitude of swelling of the triceps surae mus-

244 cles differs depending on the specific calf raise exercise. Our findings are in line with previous

245 investigations that observed differential muscle swelling and fluid shift in other muscles (e.g.,

246 quadriceps and pectoralis major) when performing different resistance exercises (1, 19, 27). The

247 first major finding of this study was the greater gastrocnemius muscle swelling observed after

248 STRA-leg than BENT-leg calf raise. Muscle swelling occurs with the shift in a fluid to the exer-

249 cised muscle (21, 23). That said, greater gastrocnemius muscle swelling in the STRA-leg calf raise

250 observed in the present study may be explained, at least in part, by the length of the gastrocnemius

251 muscles in the calf raise exercises and the associated consequences. Given the biarticular feature

252 of the gastrocnemius muscles, in the STRA-leg calf raise, both heads are likely to have a better

253 length-tension relationship. This factor may have favored the formation of an optimal number of

254 actin and myosin cross-bridges (14). Thus, the higher metabolic activity would create an aug-

255 mented demand for ATP and a greater accumulation of byproducts in active muscles (21, 23). This

256 greater byproduct accumulation might favor fluid shifts, then muscle swelling. On the other hand,

257 in the BENT-leg calf raise exercise, gastrocnemius probably experienced active insufficiency at

258 which there was little opportunity for cross-bridge cycling (14), potentially reducing the accumu-

259 lation of byproducts and, consequently, a reduced muscle swelling in gastrocnemius muscles.

260 Intriguingly, we observed a significant change in GL swelling after the BENT-leg calf

261 raise, but not in the GM head. Theoretically, GM and GL experience active insufficiency at higher

262 knee flexion angles, but our results suggest that this was not necessarily the case. Given the muscle

263 swelling observed in the lateral head, this belly likely contributed relevantly to the plantar flexion

264 task in the BENT-leg calf raise. Our results corroborate a previous study (24) in which the GL

265 does not reduce its activation during the plantar flexion task performed at 90º of knee flexion. On
266 the other hand, our findings contradict the results of another report (20) that observed no pre-to-

267 post training session changes in the transverse relaxation time of the GL at 90º of knee flexion,

268 suggesting no relevant contribution of GL at this knee angle. In regarding our results, a potential

269 explanation for muscle swelling in the GL but not in the GM may be at the origin of these muscles

270 (8, 24); the GM origin is attached closer to the midline of the knee joint and more dorsal than the

271 origin of the GL head (8). This might favor the GM to experience greater active insufficiency than

272 the GL head in BENT-leg calf raise exercise. In fact, results suggesting that the GM experience

273 active insufficiency are more consistent (4, 20, 24).

274 The second major finding of this study was greater SOL muscle swelling after BENT-leg

275 than STRA-leg calf raise. Interestingly, this greater SOL muscle swelling occurred with the abso-

276 lute load in BENT-leg being lower than in STRA-leg calf raise. A potential explanation for this

277 result may be the reduction in the participation of gastrocnemius heads with the knees bent that

278 occurred in the BENT-leg calf raise. Indeed, in the present study, the GM did not show any changes

279 in muscle thickness immediately post the BENT-leg calf raise. Although the GL showed substan-

280 tial muscle swelling, this increase was about half the swelling that occurred after the straight-legcalf

281 raise (7.0% vs. 14.5%, BENT-leg, and STRA-leg calf raise, respectively). Due to this lower partic-

282 ipation of the gastrocnemius, the plantar flexion task was performed mainly by the SOL. SOL

283 muscle originates on the soleal line of the tibia and posterior head and the upper shaft of the fibula

284 (8). Thus, changes in knee angle do not influence its length and force-generating capacity. In this

285 sense, the SOL likely exerted a greater role in the plantar flexion task in BENT-leg calf raise. As

286 a result, the higher metabolic activity would create an augmented demand for ATP, which probably

287 elicited a greater accumulation of byproducts and superior SOL muscle swelling in the BENT-leg

288 than in the STRA-leg calf raise.

289 Despite the plausibility of the argumentation proposed above, it is important to note that

290 findings on muscular activation (measured by sEMG) are inconclusive in demonstrating whether

291 there is or not greater excitation of the SOL muscle during BENT-leg calf raise task (4, 7, 24). For

292 instance, SOL sEMG amplitude during the BENT-leg was not superior to the STRA-leg exercise,

293 despite the reduction in the activation of the gastrocnemius with increased knee flexion (7). A

294 possible explanation for discordant findings on the magnitude of SOL activation (measured via

295 sEMG) and muscle swelling may be the occurrence of motor unit cycling (18, 28). During fatigu-

296 ing contractions, some motor units may shut off their firing frequencies to allow for recovery from

297 fatigue (18, 31). As a certain force is still needed to maintain the adequate force output to continue

298 exercising, these fatigued motor units will transfer their force production duties onto other motor

299 units—thus activating further muscle fibers (9, 18). This would not necessarily elicit greater sEMG

300 amplitude but would result in greater muscle swelling by activating further SOL muscle fibers;

301 particularly due to the lower gastrocnemius participation in BENT-leg calf raise. This hypothesis

302 conceivably helps to explain why there was greater SOL muscle swelling in the BENT-leg, even

303 with a lower absolute load than in STRA-leg calf raise. Importantly, in the present investigation,

304 we exclusively quantified muscle swelling, thus studies measuring motor unit recruitment, muscle

305 fiber activation (via alternative methods such as fiber glycogen depletion) and muscle swelling are

306 needed to test these hypotheses.

307 Our study has some limitations that must be acknowledged. First, we measured the change

308 in muscle thickness in one site of each muscle to express the muscle swelling; a more valid proxy

309 would be the change in whole muscle volume. Also, measures at different regions could indicate

310 whether swelling could be regional, similar to what was demonstrated with transverse relaxation

311 time after STRA-leg calf raise (15). Second, although subjects were instructed to perform the
312 exercises at the same execution velocity and range of motion, no device was used to monitor these

313 factors strictly. Third, we verified the effects of calf raise in only two knee flexion angles; future

314 investigations analyzing other knee angles may provide additional insights. Nonetheless, the two

315 exercises investigated in the present study are frequently included in resistance training routines

316 (10). In this sense, our results are of high external validity and practical relevance. Finally, we

317 measured only young women; future studies may consider testing subjects of different ages, to see

318 if this muscle ability to shift the focus of activation between muscles is preserved or altered with

319 aging, sex, or training status.

320

321 PRACTICAL APPLICATION

322 STRA-leg calf raise elicited substantial muscle swelling in the three muscles of the triceps

323 surae, while BENT-leg calf raise elicited muscle swelling in the GL and SOL, but not in the GM.

324 Notably, muscle swelling in the GM and GL was greater when performing the STRA-leg than

325 BENT-leg calf raise exercise. Conversely, SOL muscle swelling was greater when performing the

326 BENT-leg than STRA-leg calf raise. From a practical perspective, if the aim is to provide stimulus

327 to the three triceps surae muscles, the STRA-leg calf raise exercise should be preferred. If the

328 target muscle is the SOL, BENT-leg calf raise should be prioritized. Importantly, if the objective

329 is to optimally stimulate all three muscles, strength and conditioning professionals and practition-

330 ers should consider performing both calf resistance exercises. Notably, whether this pattern of

331 acute changes corresponds to hypertrophic adaptations in the gastrocnemius heads and SOL re-

332 mains to be determined.

333 REFERENCE

334 1. Albarello JCS, Cabral HV, Leitão BFM, Halmenschlager GH, Lulic-Kuryllo T, and Matta

335 TT. Non-uniform excitation of pectoralis major induced by changes in bench press

336 inclination leads to uneven variations in the cross-sectional area measured by panoramic

337 ultrasonography. J Electromyogr Kinesiol 67: 102722, 2022.

338 2. Albracht K, Arampatzis A, and Baltzopoulos V. Assessment of muscle volume and

339 physiological cross-sectional area of the human triceps surae muscle in vivo. J Biomech

340 41: 2211-2218, 2008.

341 3. Antonio J. Nonuniform response of skeletal muscle to heavy resistance training: Can

342 bodybuilders induce regional muscle hypertrophy? J Strength Cond Res 14: 102-113, 2000.

343 4. Arampatzis A, Karamanidis K, Stafilidis S, Morey-Klapsing G, DeMonte G, and

344 Brüggemann GP. Effect of different ankle- and knee-joint positions on gastrocnemius

345 medialis fascicle length and EMG activity during isometric plantar flexion. J Biomech 39:

346 1891-1902, 2006.

347 5. Cagnie B, Elliott JM, O'Leary S, D'Hooge R, Dickx N, and Danneels LA. Muscle

348 functional MRI as an imaging tool to evaluate muscle activity. J Orthop Sports Phys Ther

349 41: 896-903, 2011.

350 6. Cohen J. A power primer. Psychol Bull 112: 155-159, 1992.

351 7. Cresswell AG, Löscher WN, and Thorstensson A. Influence of gastrocnemius muscle

352 length on triceps surae torque development and electromyographic activity in man. Exp

353 Brain Res 105: 283-290, 1995.

354 8. Dalmau-Pastor M, Fargues-Polo B, Jr., Casanova-Martínez D, Jr., Vega J, and Golanó P.

355 Anatomy of the triceps surae: a pictorial essay. Foot Ankle Clin 19: 603-635, 2014.
356 9. Dankel SJ, Mattocks KT, Jessee MB, Buckner SL, Mouser JG, and Loenneke JP. Do

357 metabolites that are produced during resistance exercise enhance muscle hypertrophy? Eur

358 J Appl Physiol 117: 2125-2135, 2017.

359 10. Hackett DA. Training, supplementation, and pharmacological practices of competitive

360 male bodybuilders across training phases. J Strength Cond Res 36: 963-970, 2022.

361 11. Hackett DA, Johnson NA, Halaki M, and Chow CM. A novel scale to assess resistance-

362 exercise effort. J Sports Sci 30: 1405-1413, 2012.

363 12. Hirono T, Ikezoe T, Taniguchi M, Tanaka H, Saeki J, Yagi M, Umehara J, and Ichihashi

364 N. Relationship between muscle swelling and hypertrophy induced by resistance training.

365 J Strength Cond Res 36: 359-364, 2022.

366 13. Kassiano W, Costa B, Kunevaliki G, Soares D, Zacarias G, Manske I, Takaki Y, Ruggiero

367 MF, Stavinski N, Francsuel J, Tricoli I, Carneiro MAS, and Cyrino ES. Greater

368 gastrocnemius muscle hypertrophy after partial range of motion training performed at long

369 muscle lengths. J Strength Cond Res Epub ahead of print, 2022.

370 14. Kawakami Y, Ichinose Y, and Fukunaga T. Architectural and functional features of human

371 triceps surae muscles during contraction. J Appl Physiol (1985) 85: 398-404, 1998.

372 15. Kinugasa R, Kawakami Y, and Fukunaga T. Muscle activation and its distribution within

373 human triceps surae muscles. J Appl Physiol 99: 1149-1156, 2005.

374 16. Maeo S, Saito A, Otsuka S, Shan X, Kanehisa H, and Kawakami Y. Localization of muscle

375 damage within the quadriceps femoris induced by different types of eccentric exercises.

376 Scand J Med Sci Sports 28: 95-106, 2018.

377 17. Nunes JP, Costa BDV, Kassiano W, Kunevaliki G, Castro-e-Souza P, Rodacki ALF, Fortes

378 LS, and Cyrino ES. Different foot positioning during calf training to induce portion-

379 specific gastrocnemius muscle hypertrophy. J Strength Cond Res 34: 2347-2351, 2020.

380 18. Person RS. Rhythmic activity of a group of human motoneurones during voluntary

381 contraction of a muscle. Electroencephalogr Clin Neurophysiol 36: 585-595, 1974.

382 19. Ploutz-Snyder LL, Convertino VA, and Dudley GA. Resistance exercise-induced fluid

383 shifts: change in active muscle size and plasma volume. Am J Physiol 269: R536-543,

384 1995.

385 20. Price TB, Kamen G, Damon BM, Knight CA, Applegate B, Gore JC, Eward K, and

386 Signorile JF. Comparison of MRI with EMG to study muscle activity associated with

387 dynamic plantar flexion. Magn Reson Imaging 21: 853-861, 2003.

388 21. Raja MK, Raymer GH, Moran GR, Marsh G, and Thompson RT. Changes in tissue water

389 content measured with multiple-frequency bioimpedance and metabolism measured with

390 31P-MRS during progressive forearm exercise. J Appl Physiol (1985) 101: 1070-1075,

391 2006.

392 22. Schoenfeld BJ. Accentuating muscular development through active insufficiency and

393 passive tension. Strength Cond J 24: 20-22, 2002.

394 23. Schoenfeld BJ. Potential mechanisms for a role of metabolic stress in hypertrophic

395 adaptations to resistance training. Sports Med 43: 179-194, 2013.

396 24. Signorile JF, Applegate B, Duque M, Cole N, and Zink A. Selective recruitment of the

397 triceps surae muscles with changes in knee angle. J Strength Cond Res 16: 433-439, 2002.

398 25. Steele J, Fisher J, Giessing J, and Gentil P. Clarity in reporting terminology and definitions

399 of set endpoints in resistance training. Muscle Nerve 56: 368-374, 2017.
400 26. Vieira A, Blazevich A, Souza N, Celes R, Alex S, Tufano JJ, and Bottaro M. Acute changes

401 in muscle thickness and pennation angle in response to work-matched concentric and

402 eccentric isokinetic exercise. Appl Physiol Nutr Metab 43: 1069-1074, 2018.

403 27. Vieira DCL, Dourado MAA, Ugliara L, Durigan JLQ, Schoenfeld BJ, and Bottaro M. Can

404 hip joint position affect quadriceps muscle responses during knee extension exercise? Int

405 J Sports Med 41: 929-935, 2020.

406 28. Vigotsky AD, Beardsley C, Contreras B, Steele J, Ogborn D, and Phillips SM. Greater

407 electromyographic responses do not imply greater motor unit recruitment and 'hypertrophic

408 potential' cannot be inferred. J Strength Cond Res 31: e1-e4, 2017.

409 29. Vigotsky AD, Halperin I, Trajano GS, and Vieira TM. Longing for a longitudinal proxy:

410 acutely measured surface EMG amplitude is not a validated predictor of muscle

411 hypertrophy. Sports Med 52: 193-199, 2022.

412 30. Wakahara T, Fukutani A, Kawakami Y, and Yanai T. Nonuniform muscle hypertrophy: its

413 relation to muscle activation in training session. Med Sci Sports Exerc 45: 2158-2165,

414 2013.

415 31. Westad C, Westgaard RH, and De Luca CJ. Motor unit recruitment and derecruitment

416 induced by brief increase in contraction amplitude of the human trapezius muscle. J Physiol

417 552: 645-656, 2003.

418
419

420 Figure 1. Representation of the ultrasound sites in the GM, GL, and SOL muscles (A). The ultra-

421 sonography images show GM, GL, and SOL muscles (B). For each muscle, superficial and deep

422 aponeuroses are visualized, between which the muscle thickness was measured. GM = gas-

423 trocnemius medial; GL = gastrocnemius lateral; SOL = soleus.

Figure 2. STRA-leg (A) and BENT-leg calf raise (B) exercises. STRA-leg = straight-leg calf raise,

BENT-leg = bent-leg calf raise.

Table 1. Repetitions performance and RIR in the two experimental training sessions (n = 17).
STRA-leg BENT-leg
1st set 2nd set 3rt set 4th set 1st set 2nd set 3rt set 4th set
Repetitions (n°) 19.8 ± 0.7 18.6 ± 1.7 15.7 ± 1.9*‡ 14.6 ± 1.6*‡ 19.8 ± 0.5 18.2 ± 1.9 15.0 ± 3.3*‡ 13.8 ± 2.9*‡
RIR (n°) 0.7 ± 0.5 0.1 ± 0.3* 0.0 ± 0.2* 0.0 ± 0.2* 0.7 ± 1.1 0.1 ± 0.3* 0.0 ± 0.0* 0.0 ± 0.0*
Note. †P < 0.05 vs. BENT-leg; *P < 0.05 vs. 1st set; ‡P < 0.05 vs. 2nd set. RIR = repetitions in reserve; STRA-leg = straight-leg calf raise; BENT-leg
= bent-leg calf raise.
Table 2. Muscle thickness (cm) of the GM, GL, and SOL before and im-
mediately after STRA-leg and BENT-leg calf raise exercises (n = 17).
Muscle thickness STRA-leg BENT-leg
GM
Pre 1.90 ± 0.33 1.90 ± 0.33
Post 2.06 ± 0.35*† 1.85 ± 0.32
Meandiff 0.16 (0.12, 0.21) -0.02 (-0.05, 0.01)
ES 0.48 -0.15
GL
Pre 1.40 ± 0.31 1.38 ± 0.32
Post 1.59 ± 0.32*† 1.46 ± 0.31*
Meandiff 0.19 (0.13, 0.26) 0.09 (0.05, 0.12)
ES 0.60 0.25
SOL
Pre 2.20 ± 0.31 2.18 ± 0.31
Post 2.37 ± 0.33*† 2.51 ± 0.32*
Meandiff 0.17 (0.13, 0.21) 0.33 (0.28, 0.37)
ES 0.55 1.06
Notes. Pre- and post-training sessions data are presented as mean and
standard deviation, while meandiff as mean and 95% confidence intervals.
ES = effect size; *P < 0.05 vs. pre; †P < 0.05 vs. BENT-leg. STRA-leg =
straight-leg calf raise; BENT-leg = bent-leg calf raise.
 Figure 3. Pre-to-post changes in muscle thickness of GM, GL, and SOL in response to STRA-leg and BENT-leg calf raise exercises. Bars are

means, and circles are individual values. *P < 0.05 vs. pre; †P < 0.05 vs. BENT-leg calf raise. GM = gastrocnemius medial; GL = gastrocnem-

ius lateral; SOL = soleus; STRA-leg = straight-leg calf raise; BENT-leg = bent-leg calf raise.

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