fphys-10-01456 November 23, 2019 Time: 16:4 # 1 ORIGINAL RESEARCH published: 26 November 2019 doi: 10.3389/fphys.2019.01456 Triceps Surae Muscle Architecture Adaptations to Eccentric Training Jeam Marcel Geremia1*, Bruno Manfredini Baroni2, Rodrigo Rico Bini3, Fabio Juner Lanferdini4, Amanda Rodrigues de Lima1, Walter Herzog5 and Marco Aurélio Vaz1 1 Laboratório de Pesquisa do Exercício, Escola de Educação Física, Fisioterapia e Dança, Universidade Federal do Rio Grande do Sul, Porto Alegre, Brazil, 2 Departamento de Fisioterapia, Universidade Federal de Ciências da Saúde de Porto Alegre, Porto Alegre, Brazil, 3 Holsworth Research Initiative, La Trobe Rural Health School, La Trobe University, Bendigo, VIC, Australia, 4 Laboratório de Biomecânica, Centro de Desportos, Universidade Federal de Santa Catarina, Florianópolis, Brazil, 5 Faculty of Kinesiology, Engineering, Medicine and Veterinary Medicine, University of Calgary, Calgary, AB, Canada Background: Eccentric exercises have been used in physical training, injury prevention, and rehabilitation programs. The systematic use of eccentric training promotes specific morphological adaptations on skeletal muscles. However, synergistic muscles, such as the triceps surae components, might display different structural adaptations due to differences in architecture, function, and load sharing. Therefore, the purpose of this study was to determine the effects of an eccentric training program on the Edited by: triceps surae (GM, gastrocnemius medialis; GL, gastrocnemius lateralis; and SO, soleus) Adamantios Arampatzis, muscle architecture. Humboldt University of Berlin, Germany Methods: Twenty healthy male subjects (26 ± 4 years) underwent a 4-week control Reviewed by: period followed by a 12-week eccentric training program. Muscle architecture [fascicle Maria Elissavet Nikolaidou, National and Kapodistrian University length (FL), pennation angle (PA), and muscle thickness (MT)] of GM, GL, and SO was of Athens, Greece evaluated every 4 weeks by ultrasonography. Simon Steib, Technical University of Munich, Results: Fascicle lengths (GM: 13.2%; GL: 8.8%; SO: 21%) and ML increased (GM: Germany 14.9%; GL: 15.3%; SO: 19.1%) from pre- to post-training, whereas PAs remained *Correspondence: similar. GM and SO FL and MT increased up to the 8th training week, whereas GL, Jeam Marcel Geremia [email protected] FL increased up to the 4th week. SO displayed the highest, and GL the smallest gains in FL post-training. Specialty section: This article was submitted to Conclusion: All three synergistic plantar flexor muscles increased FL and MT with Exercise Physiology, eccentric training. MT increased similarly among the synergistic muscles, while the a section of the journal Frontiers in Physiology muscle with the shortest FL at baseline (SO) showed the greatest increase in FL. Received: 22 September 2019 Keywords: eccentric exercise, muscle architecture, muscle plasticity, triceps surae, ultrasonography Accepted: 11 November 2019 Published: 26 November 2019 Citation: INTRODUCTION Geremia JM, Baroni BM, Bini RR, Lanferdini FJ, de Lima AR, Herzog W Muscle architecture (i.e., the geometrical arrangement by which muscle fibers are organized with and Vaz MA (2019) Triceps Surae Muscle Architecture Adaptations respect to a muscle’s line of action) has an important role in skeletal muscle force production to Eccentric Training. (Fukunaga et al., 1997; Lieber and Fridén, 2000). Fascicle length (FL) is associated with the serial Front. Physiol. 10:1456. sarcomere number, which has a direct impact on the fiber/muscle shortening velocity and excursion doi: 10.3389/fphys.2019.01456 (Edman et al., 1985; Lieber and Fridén, 2000; Blazevich, 2006). Longer FL is associated with Frontiers in Physiology| www.frontiersin.org 1 November 2019| Volume 10| Article 1456 fphys-10-01456 November 23, 2019 Time: 16:4 # 2 Geremia et al. Plantar Flexor Adaptation to Eccentric Training improved performance in activities demanding high velocities on the triceps surae muscles are conflicting. While some studies of shortening (Abe et al., 2000, 2001; Kumagai et al., 2000), show that eccentric training promotes an increase in FL, PA, whereas a reduction in FL with aging explains a large part of and MT (Duclay et al., 2009; Geremia et al., 2018b), other the reduced maximal shortening velocity in older compared to studies did not find changes in these architectural outcomes young adults (Thom et al., 2007). Muscle fiber pennation angle (Raj et al., 2012; Fouré et al., 2013). In the majority of studies (PA) is related to the parallel number of sarcomeres within a fiber, (Duclay et al., 2009; Raj et al., 2012), it was assumed that and is related to the fiber diameter and its maximal capacity for adaptations observed for the gastrocnemius medialis (GM) were force production (Fukunaga et al., 1997; Blazevich and Sharp, representative of the entire triceps surae muscle group. However, 2005). Hypertrophied muscles (i.e., with a large number of different muscle architecture adaptations were observed after sarcomeres in parallel) tend to have high PA (Kawakami et al., 6 weeks of stretching between the GM and the gastrocnemius 1993), while muscles of elderly (Narici et al., 2003) or disuse lateralis (GL) (Simpson et al., 2017). This may be due to atrophied muscles (Suetta et al., 2009) (i.e., with a small number the fact that load sharing between the synergistic muscles is of parallel sarcomeres), tend to have small PA. Muscle thickness not homogeneous, and different mechanical loads have been (MT) is influenced by both the serial and parallel sarcomere observed for the different components of the triceps surae number, as it depends on FL and PA (Fukunaga et al., 1997; muscle (Crouzier et al., 2018). Also, differential adaptations to Baroni et al., 2013b). eccentric training have been observed for the vastus lateralis Structural changes accompanying (or due to) strength training and vastus intermedius of rats exposed to a chronic downhill provide insights into the muscle’s ability to accommodate training protocol (Butterfield et al., 2005). Furthermore, short architecture adaptations specific to the movement’s mechanical muscle fibers are more susceptible to muscle damage caused by demands. Although traditional isoinertial resistance training is eccentric training than long muscle fibers (Proske and Morgan, executed with a constant external load in the concentric and 2001; Baroni et al., 2013b). Considering the differences in muscle eccentric phases of movement (Walker et al., 2016), training fascicular geometry of the triceps surae components (GM, GL, programs often involve isometric contractions (Kitai and Sale, and SO, soleus) (Wickiewicz et al., 1983; Maganaris et al., 1998; 1989; Oranchuk et al., 2019) or have an emphasis on the Fouré et al., 2013), eccentric exercise may cause different levels of concentric (Blazevich et al., 2007; Timmins et al., 2016b) or micro-damage in these muscles. Therefore, the idea of a specific eccentric (Baroni et al., 2013b; Geremia et al., 2018b) phases. adaptive response for the different components of the triceps Evidence suggests that training programs neglecting eccentric surae muscle seems reasonable. actions do not prepare subjects for the eccentric demands We have previously demonstrated time-dependent encountered in sports and activities of daily living (Barstow et al., architectural adaptations in human knee extensor muscles 2003; LaStayo et al., 2003; Lovering and Brooks, 2014; Franchi (Baroni et al., 2013b). However, we were unable to find studies et al., 2017). Therefore, strength training with an emphasis evaluating time-dependent adaptations in muscle architecture on eccentric movement execution has become popular for the for the three triceps surae muscles. Considering the conflicting purpose of injury prevention (LaStayo et al., 2003; Goode results regarding eccentric training adaptations of the triceps et al., 2015) and rehabilitation (Rees et al., 2008; Murtaugh surae muscles, the limited information on architectural and Ihm, 2013; Frizziero et al., 2014), as well as physical adaptations of the individual triceps surae muscles, and the lack fitness improvement in healthy subjects (Baroni et al., 2013c; of longitudinal, time-dependent adaptations of muscles subjected Geremia et al., 2018a,b). to eccentric training programs, the purpose of this study was to Several studies demonstrated changes in knee extensor determine the effects of a 12-week eccentric training program on muscle architecture with eccentric training (Higbie et al., 1996; the architecture of the GM, GL, and SO muscles. Based on the Hortobágyi et al., 1996; Housh et al., 1996; Seger et al., 1998; literature (Proske and Morgan, 2001; Duclay et al., 2009; Baroni Reeves et al., 2003; Blazevich et al., 2007; Santos et al., 2010; Raj et al., 2013b; Geremia et al., 2018b), we hypothesize that eccentric et al., 2012; Baroni et al., 2013b,c, 2015). Most of these studies training will promote adaptations in muscle architecture, causing reported an increase in FL (Blazevich and Sharp, 2005; Blazevich larger increases in FL and MT in the muscle with the shortest FL et al., 2007; Baroni et al., 2013b), while results on changes in PA (i.e., SO). In addition, we also expected that these adaptations were mixed (Blazevich et al., 2007; Reeves et al., 2009; Raj et al., will occur between 4 and 8 weeks of the training program. 2012; Baroni et al., 2013b). Eccentric training has been found to produce significant changes in muscle architecture after training periods