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See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/362864671 Muscle Hypertrophy Response to Range of Motion in Strength Training: A Novel Approach to Understanding the Findings Article in Strength & Conditioning Journal · August 2022 DOI: 10.1519/SSC.0000000000000737 CITATIONS 15 READS 10,835 6 authors, including: Charlie Ottinger Barton College 15 PUBLICATIONS 55 CITATIONS SEE PROFILE Matthew H Sharp 47 PUBLICATIONS 374 CITATIONS SEE PROFILE Matthew W Stefan 26 PUBLICATIONS 107 CITATIONS SEE PROFILE Raad H Gheith Applied Science and Performance Institute 17 PUBLICATIONS 78 CITATIONS SEE PROFILE All content following this page was uploaded by Charlie Ottinger on 17 May 2023. 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Ottinger, Matthew H. Sharp, Matthew W. Stefan, Raad H. Gheith, Fernando de la Espriella, and Jacob M. Wilson Research Division, Applied Science and Performance Institute, Tampa, Florida A B S T R A C T One resistance training variable that may be altered to achieve desired outcomes is the range of motion used in training. Generally, the strength and conditioning field has accepted that using a greater range of motion in strength training exercises results in more substantial muscle hypertrophy outcomes. How- ever, this theory has proved to be inconsistently supported in the literature, and to date, no sufficient explanation exists to explain this phenomenon. This review article seeks to outline a novel approach for potentially describing the disparities seen in range of motion research with respect to hypertrophy outcomes by applying the unique length- tension curve of each muscle being examined. As will be discussed in the review, virtually all the results from range of motion studies in various muscles have corresponded to each muscle’s length-tension curve; muscles that are active on the descending limb of the curve appear to garner greater hyper- trophy from using larger ranges of motion. Conversely, muscles that are not active on the descending limb exhibit similar adaptations despite alterations in range of motion. A novel hypothesis for applying this information to resistance training programs will be presented and discussed. INTRODUCTION D esigning safe and effective strength and conditioning pro- grams requires careful and purposeful institution of appropriate training variables for the desired out- come. Given that resistance training adaptations are specific to the mode or variable of training performed (20), it is necessary to implement these variables in a planned fashion specific to the goals of the athlete or team performing the program. Common training variables used in resistance training programs include repetition ranges (61), training intensity (39,61), training frequency (63), contraction speed emphasized (52,62), loading pat- terns (33,70), and even the joint range of motion used (5,19,32,38,55,59,68) in resistance training exercises. The range of motion used during resis- tance training exercises is a unique train- ing variable in that it affects long-term adaptations but is also influenced by the inherent biomechanics and abilities of the trainee. Indeed, unique individual anthropometry largely determines the mechanical challenge imposed by resis- tance training exercises because these Address correspondence to Charlie R. Ot- tinger, cottinger@theaspi.com. KEY WORDS : range of motion; length-tension curve; hypertrophy; muscle length Copyright � National Strength and Conditioning Association Strength and Conditioning Journal | www.nsca-scj.com 1 Copyright © National Strength and Conditioning Association. Unauthorized reproduction of this article is prohibited. mailto:cottinger@theaspi.com inherent kinesthetic traits largelydictate the overall range of motion performed at each joint involved in an exercise (9). In addition, training experience in both stretching and resistance training can improve joint flexibility (41,65) which leads to an improved ability to withstand a greater range of motion in strength training exercises. Finally, the intrinsic or trained flexibility of a given muscle can also influence its optimal length for force production, thus adjusting the length-tension curve of the muscle (75). In applied settings, this affects exercise selection and/or modification as well as the ranges of motion used. Researchers can measure strength training range of motion through a small variety of tools. The primary mea- surement method has centered on assess- ing the degrees of rotation performed at a specific joint (5,19,32,38,55,68). In addi- tion, resistance implement displacement has also been used in research (26) albeit to a lesser extent. Ultimately, joint range of motion is likely the most common method of measurement because this cal- culation is more relevant to the training stimulus a givenmuscle may receive in an exercise. This is due to the relationship between the range of motion performed at a joint and the concomitant length change of each muscle that interacts with the joint (51). Indeed, previous studies have consis- tently used greater ranges of motion or deeper positions of joint flexion to assess muscles at “long” lengths, whereas short- er ranges of motion or more shallow joint angles represent “short” muscle lengths (19,37,38,47,55,57,59). Certainly, the change in joint angle does not per- fectly correlate with a change in muscle length because this relationship can be affected by muscle activation (48,64), contraction/movement velocity (43), and muscle pennation angle and resul- tant architectural gear ratios (64). How- ever, the pursuit of ecological validity reigns supreme in the field of applied strength and conditioning, and joint range of motion remains the primary way of dictating training with “long” or “short” muscle lengths. Because ultraso- nography is necessary to truly measure muscle length (25,68), this tool is rarely used in strength and conditioning set- tings. One important distinction is that the specific joint degrees used in various ranges of motion are not interchange- able. Indeed, moving from 0 to 508 of elbow flexion involves the same absolute range of motion as 50–1008 elbow flex- ion; however, the 0–508 range would certainly be considered a longer length than the 50–1008 range. Regardless, in many applied strength and conditioning studies, the terms “range of motion” and “muscle length” are almost used inter- changeably in which greater ranges of motion refer to greater muscle lengths and vice versa. With this application in mind, Bloomquist et al. (5) and McMahon et al. (38) found that greater ranges of motion at the knee were more effective at inducing muscle hypertrophy in the quadriceps muscle group; however, Pinto et al. (55) discov- ered that partial range of motion elbow flexion training was just as effective at pro- moting muscle hypertrophy in the biceps as full range of motion training. It is gen- erally accepted that full range of motion training is likely optimal for muscular development, but this paradigm has been met with contrary findings from Pinto et al. (55) and both Goto et al. (19) and Stasinaki et al. (68) who found similar lev- els of triceps hypertrophy after training with full or partial elbow extension ranges of motion. Furthering the confusion, recent research from Maeo et al. (37) uncovered that seated knee flexion train- ing produced greater hypertrophic adap- tations in the hamstrings compared with prone knee flexion training. Because the hamstrings are a biarticulate muscle group that crosses both hip and knee joint, the seated knee flexion variation lengthens the hamstrings to a greater degree (36), thus imposing a greater range ofmotion for the hamstrings. Although the aforementioned research into the range of motion used during resistance training exercises has pro- duced unclear results, the mechanism through which these adaptations take place may not be as simple as previously believed. The weight of the research has begun to show that different muscle groups adapt uniquely to different ranges of motion which warrants further theo- retical investigation to explain this phe- nomenon. Therefore, the purposes of this article were to (a) review the process of muscle hypertrophy and how muscle length can affect hypertrophic adapta- tions, (b) analyze previous findings related to range of motion and muscle hypertrophy, and (c) introduce a novel theory for explaining these occasionally contrary findings. MUSCLE HYPERTROPHY Because muscle hypertrophy is a pri- mary outcome in many range of motion studies, a brief review of this phenomenon is necessary. Muscle hypertrophy is the procedure through which a muscle fiber undergoes a re- modeling process which, in turn, results in a larger muscle fiber. Muscle hypertrophy can occur through at least 3 unique morphological pro- cesses, including sarcoplasmic hyper- trophy, myofibrillar hypertrophy, and longitudinal hypertrophy. Sarcoplasmic hypertrophy is typically referred to as an increase in cellular sar- coplasm or sarcoplasmic components which results in the expansion of the overall volume of the muscle fiber (22,58). Previously, it was believed that endurance or interval training had a greater association with sarcoplasmic hypertrophy because of an increase in muscle glycogen storage; however, recent research from Haun et al. (21) uncovered that resistance training can also induce sarcoplasmic hypertrophy, even to a disproportionate ratewithmyo- fibrillar hypertrophy which is more often associated with resistance training. Although the exact mechanism that pref- erentially induces sarcoplasmic hypertro- phy over myofibrillar hypertrophy is unclear, sarcoplasmic hypertrophy plays at least a minor role in hypertrophic adaptations to resistance training (58). Myofibrillar hypertrophy is the most common type of muscle hypertrophy associated with resistance training. This is seen in both acute studies in which myofibrillar protein synthesis rates increase after resistance training Muscle Hypertrophy Response VOLUME 00 | NUMBER 00 | MONTH 20222 Copyright © National Strength and Conditioning Association. Unauthorized reproduction of this article is prohibited. (13) and longitudinal studies in which trained subjects exhibit higher levels of myofibrillar proteins than untrained individuals (72). Myofibrillar hypertro- phy occurs through an increase in myofilament packing density, or more simply put, an increase in the abun- dance of contractile proteins within the myofibril (58). Ultimately, this structural adaptation increases the overall diameter of the muscle fiber. It is posited that myofibrillar hypertro- phy is mostly associated with resis- tance training because it is highly related to the force producing ability of a muscle (69). Therefore, the most prominent mechanism through which myofibrillar hypertrophy occurs is likely resistance training. Longitudinal hypertrophy is displayed through a lengthening of the myofibril. Previously, researchers understood this change in length to be a result of an increase of sarcomeres in series, which would effectively lengthen the muscle fiber (15). However, more recent research has highlighted that an increase of sarcomeres in series is not necessary to elicit longitudinal hyper- trophy (54), so further research is cer- tainly needed to further understand this form of muscle hypertrophy. Regardless, this unique adaptation is believed to be preferentially induced during eccentric contractions or even active and passive stretching in which a muscle experiences tension at long muscle lengths. Mechanistically, longi- tudinal hypertrophy may be a protec- tive adaptation to better produce force at long muscle lengths (16,40); how- ever, furtherresearch is needed to ver- ify this theory. Moreover, the visual microscopic representation of longitu- dinal hypertrophy can be difficult to quantify (29), thus increasing the need for further exploration into this topic. LENGTH-TENSION CURVE AND RESISTANCE TRAINING APPLICATIONS LENGTH-TENSION CURVE Although the relationship between potential hypertrophic stimuli and observed muscle hypertrophy is not fully understood, it is well accepted that mechanical tension is likely the primary driver of myofibrillar and lon- gitudinal hypertrophy (74). Mechani- cal tension is detected by mechanoreceptors within the muscle that, in essence, measure muscular force production through deformation of their plasma membrane. After detec- tion of sufficient force, mechanorecep- tors create electrical and chemical signals that begin the protein remodel- ing process that results in muscular hypertrophy (6,74). However, the route of mechanical tension detection that stimulates longitudinal hypertro- phy may be unique. In 1954, Huxley and Niedergerke (27) published the first ever report that detailed findings related to microscopic analysis of living muscle fibers. This article was shortly followed by a similar article from Huxley and Hanson (28). The combination of these articles led to the birth of the sliding filament the- ory, which suggested that muscles con- tract using 2 proteins, actin and myosin, “sliding” past one another. However, this theory did not explain every aspect of muscular contraction; factors such as residual force enhancement and passive force production were difficult to explain through this model (34). More recent research has proposed an update to the sliding filament theory: the wind- ing filament theory. The winding fila- ment theory builds on the sliding filament theory but fills the gaps through the inclusion of the giant pro- tein, titin. As the theory’s name sug- gests, titin “winds” itself around the contractile protein, actin, during length- ening contractions and then releases stored energy like a spring during short- ening contractions (16,45,46,66). Intriguingly, titin in and of itself is a type of mechanoreceptor that detects mechanical tension. Owing to the spring-like action of titin, various aspects of its winding and unfolding process sense mechanical loading much like the aforementioned mechanoreceptor (31). Titin’s mechanosignaling process differs from mechanoreceptors through unique pathways and downstream signaling molecules, but the end process of this signaling cascade results in protein remodeling that ultimately leads to mus- cle hypertrophy (35). A second distinc- tive factor of titin is that its resultant signaling cascade is likely greatest during muscle contractions performed in a stretched or lengthened position (42). This unique aspect of titin has been the- orized to be a determinant of the greater hypertrophic adaptations occasionally seen with longer range of motion train- ing (31) because these ranges will impose greater length changes on the muscle. However, this theory is highly related to the next topic worthy of discussion: the length-tension curve. The length-tension curve visualizes the relationship between tension (or force) and sarcomere length (36) as shown in Figure 1. Sarcomeres are the individual contractile units within a single myofi- bril; muscle lengthening or shortening is dictated by the collective change of length of the sarcomeres within a sin- gle fiber. Practitioners often use this length-tension curve in tandem with the working range of motion of the primary joint a muscle acts on (56). The length-tension curve contains 3 unique components: the ascending limb, the plateau portion, and the descending limb. The plateau portion of the length- tension curve is associated with sarco- mere lengths near the resting value. The ascending limb, then, represents shorter sarcomere lengths, whereas the descending limb depicts longer sarco- mere lengths. As shown in Figure 1, the ascending and descending limbs of the length-tension curve are generally associated with increasing and decreas- ing force production, respectively. How- ever, the plateau portion of the length- tension curve displays the optimal force producing length for a muscle (36). Therefore, the astute practitioner would theoretically be correct in assuming that training a muscle near its resting length would be optimal for muscle hypertro- phy because of peak force production occurring near this position. However, the simple length-tension curve does not account for passive Strength and Conditioning Journal | www.nsca-scj.com 3 Copyright © National Strength and Conditioning Association. Unauthorized reproduction of this article is prohibited. elements, such as titin, that can also contribute to muscle force production. Thus, an adjusted length-tension curve is shown in Figure 2 in which passive elements, such as titin, muscle fascia, and tendons, also contribute to the total tension developed by the mus- cle (24). A perusal of the adjusted length- tension curve now allows the practi- tioner to theorize that long sarcomere lengths are associated with the highest muscle forces because the combination in active and passive force likely sur- passes the peak of the plateau region (24). This unique finding has likely driven the strength and conditioning field to generally accept that a larger range of motion is more effective for optimal muscle development than a shorter range because a large joint range of motion will induce the great- est overall length change in the mus- cles acting on the joint (50,56). The practice of continually maximizing ten- sion through using a full range of motion, then, should lead to greater hypertrophic gains over a long-term training program (5,32,38). However, a proverbial speed bump appears when attempting to generalize this theory to all the major muscle groups used in sport or fitness training. Intriguingly, not every muscle is active on all portions of the length-tension curve. Therefore, training adaptations to differing ranges of motion and asso- ciated muscle lengths are likely unique to the muscle group being trained. This can explain the discrepancy in findings between Bloomquist et al. (5) and Pinto et al. (55). Bloomquist et al. (5) found that deep squat training (0– 1208 knee flexion) led to greater hypertrophy in the quadriceps muscle group (4–7% increase in CSA vs. 0%) than shallow squat training (0–608 knee flexion), whereas Pinto et al. (55) found no significant difference in biceps hypertrophy (9.65 6 4.4% increase in muscle thickness vs 7.83 6 4.9%) after training programs with either a full (0– 1308 elbow flexion) or partial (50–1008 elbow flexion) range of motion, respectively. Although these findings at first seem contradictory to the general theory of full range of motion training, they may be explained rather simply. The quadriceps is primarily active on the descending limb of the length- tension curve (67) as shown in Figure 3. Therefore, using a greater range of motion at the knee during quadriceps exercises will likely result in greater hypertrophic adaptations because this muscle group experiences greater ten- sion at longer lengths. However, the biceps brachii is only active on the ascending portion of the curve (30) as shown in Figure 4. Therefore, the biceps is not active on the descending portion of the length-tension curve and receives no additional tension-related benefits from developing force at longer muscle lengths. Owing to the potential relationship between the length-tension curve and adaptations to differing ranges of motion in strength training, it is worth reviewing how altering the range of motion used in an exercise has been shown to affect long-term hypertro- phic adaptations in a variety of muscle groups. QUADRICEPS Kubo et al. (32) examined 17 young men (20.76 0.4 years) performing deep squat (0–1408 knee flexion) or half squat (0–908 knee flexion) training for 10 weeks. Both groups trained2x/week during this period and followed a linear program in which training intensity increased each week. After the 10-week training program, the authors usedMRI technology to assess the change in quadriceps, adductor, hamstrings, and Figure 1. Length-tension curve. From Lieber et al. (36). Figure 2. Adjusted length-tension curve. From Herzog (24), with permission. Muscle Hypertrophy Response VOLUME 00 | NUMBER 00 | MONTH 20224 Copyright © National Strength and Conditioning Association. Unauthorized reproduction of this article is prohibited. gluteus maximus cross-sectional area (CSA). Intriguingly, increases in quadri- ceps CSA were similar between groups (4.9 6 2.6% in full squats vs 4.6 6 3.1% in half squats), whereas increases in adductor CSAwere significantly greater in the full squat group than the half squat group (6.2 6 2.6% vs. 2.7 6 3.1%; p 5 0.026) as were changes in gluteus CSA (6.7 6 3.5% vs 2.2 6 2.6%; p 5 0.008). At a glance, (32) the findings of Kubo et al. regarding quadriceps hypertrophy in half squats seem at odds with the results from Bloomquist et al. (5) in which partial squats were shown to not induce hypertrophy in the quadri- ceps. Both studies used untrained sub- jects; however, Bloomquist et al. (5) had participants squat to a knee flexion angle of 608, whereas Kubo et al. (32) used a knee flexion angle of 908 in the half squat group. In addition, the study by McMahon et al. (38), discussed in further detail below, used a knee angle of 508 in short range of motion training and found that this range of motion was less effective for promoting quadriceps hypertrophy than using a knee angle of 908. Because the quadriceps is primarily active on the descending portion of the length-tension curve (67), it would appear that 908 of knee flexion is suffi- cient for increasing total tension and, therefore, hypertrophy in the quadriceps; however, further investigation is certainly warranted to confirm this theory. McMahon et al. (38) used 26 untrained and healthy subjects aged 19.0 6 3.4 years in an investigation seeking the effects of full training programs involv- ing either a long range of motion (0– 908 knee flexion) or a short range of motion (0–508 knee flexion) quadriceps exercises. Participants performed their respective training programs 3x$wk21 for 8 weeks, and muscle CSA of the quadriceps was measured before and after the training program at 25, 50, and 75% of femur length. The long range of motion group exhibited sig- nificantly greater increases in vastus lateralis CSA at 75% femur length as opposed to the short range of motion group (59 6 15% vs 16 6 10%; p , 0.05) after the 8-week training pro- tocol. Intriguingly, both groups made similar (p . 0.05 between groups) increases in vastus lateralis CSA at 25% (33.83 vs. 19.04% long range vs. short range, respectively) and 50% femur length (18.00% for both groups). In addition, muscle fascicle length was measured at 25, 50, and 75% of femur length. Significant differences between groups were found for fascicle lengths measured at 50% femur length (23 6 5% vs. 10 6 2%; p , 0.05) and 75% femur length (19 6 3% vs. 11 6 2; p , 0.05), both of which favored the long range of motion training group. Owing to the quadriceps position on the de- scending region of the length-tension curve, these results are not surprising. Moreover, these data may present fur- ther evidence for longitudinal hypertro- phy because both training groups exhibited significant increases in muscle fascicle length. However, the long range of motion training group experienced even greater changes in fascicle length than the short range of motion group. Another more current study per- formed by Pedrosa et al. (53) sought to examine the effects of multiple ranges of motion in the knee extension exercise on quadriceps adaptations. The authors used 45 untrained women and split these subjects into 4 groups: a full ROM (100-308 knee flexion), an initial partial ROM (100-658), a final partial ROM (65-308), and a varied ROM which alternated the initial and final partial ROM conditions. Similar to other investigations, rectus femoris (RF) and vastus lateralis (VL) muscle hypertrophy adaptations were mea- sured at 70, 60, 50, and 40% femur length. Intriguingly, the initial ROM group exhibited the greatest RF and VL hypertrophy at 70% femur length (32% & 19%, respectively, vs. 24% & 1 and 19% & 12% for final partial and full ROM, respectively) and presented with greater hypertrophy at 60% (24% & 15 vs. 6% & 5.5 and 18% & 13%) and 50% (28% & 16 vs. 13% & 11 and 13% & 14%) of femur length relative to the final partial ROM and full ROM groups. An even more interesting note is that the final partial ROM group dis- played similar changes to a nontraining control group about RF and VL hyper- trophy at 60% (6% & 5.5 vs. 2% & 2%) and 70% (24% & 1 vs. 1% & 2%) femur length. This study further emboldens the theory that the quadriceps muscle group responds more favorably to training at long muscle lengths when hypertrophy is the desired training out- come. Again, this theory is likely dependent on the length-tension curve Figure 3. Length-tension relationship of the vastus lateralis. Adapted from Son et al. (67) with permission. Strength and Conditioning Journal | www.nsca-scj.com 5 Copyright © National Strength and Conditioning Association. Unauthorized reproduction of this article is prohibited. of the quadriceps which favors increased tension at longer muscle lengths. TRICEPS Investigations from Goto et al. (19) and Stasinaki et al. (68) found differing results from Bloomquist et al. (5), McMahon et al. (38), Kubo et al. (31), and Pedrosa et al. (53) when as- sessing ranges of motion used in train- ing the triceps brachii. Goto et al. (19) split 44 resistance trained men aged 21.6 6 1.3 years into 2 groups, one of which performed partial (45–908 elbow flexion) elbow extension training and the other performed full (0–1208 elbow flexion) elbow extension training. Each group trained their respective range of motion 3x/week for 8 weeks while following a protocol of 3 sets of 8 at their 8RM load. Curiously, triceps CSA increased to a greater extent (p , 0.05) after partial range of motion training (48.76 14.5%) as opposed to full range of motion training (28.2 6 10.9%). Goto et al. (19) theorized that the greater hypoxia experienced during partial range of motion training may be the mechanism at work because the authors reported a correlation of r 5 0.70 between an increase in CSA and time spent under the oxygen- hemoglobin dissociation curve. Intrigu- ingly, muscle activation was also higher in the partial range of motion condition (p , 0.05 between groups) which could certainly have affected hypertro- phic adaptations. However, these components may not be the only mechanisms influencing this outcome. Stasinaki et al. (68) also examined unique triceps training methods, albeit in a group of 9 untrained women aged 19.3 6 0.4 years. Each participant trained one arm with a long muscle length range of motion (70–1508 elbow flexion) and the other with a short range of motion (10–908 elbow flexion). Participants performed the training protocol 2x/week for 6 weeks, and both protocols involved perform- ing 6 sets of 6 repetitions at 85% 1RM in the assigned range of motion. After the training period, both arms experi- enced significant, but not different (p5 0.618), changes in muscle thickness in which the long length arm increased by 13.7 6 9.0% and the short length arm increased by 10.7 6 15.3%. A key aspect of this particular study is that ultrasonography was also used to val- idate the ranges of motion used with respect to muscle length of the long head of the triceps. In the long muscle length condition, subjects performed overhead triceps extensions which produced a range of fascicle lengths from 60.1 6 7.7 to 66.5 6 4.8 mm. Conversely, the short range of motion training condition generated a range of fascicle lengths from 44.2 6 5.9 to 51.3 6 4.8 mm. Fasciclelength was deter- mined to be statistically different (p , 0.05) between groups which certainly aids in the theoretical applicability of this study’s findings. Ultimately, the length-tension curve still seems to apply in both the Goto et al. (19) and Stasinaki et al. (68) stud- ies. Previous research has found that the triceps is primarily active in the plateau region of the length-tension curve (44). Therefore, the triceps pro- duces similar levels of force regardless of muscle length and, thus, would the- oretically experience similar hypertro- phic adaptations in either short or long ranges of motion. GLUTEUS Recent research from Barbalho et al. (3) sought to examine the hypertrophic adaptations of the gluteus maximus after either deep squat (0–1408 knee/ hip flexion) training or barbell hip thrust training (90-08 hip flexion) which inherently includes less hip and knee range of motion compared with the deep squat. The authors split 22 trained women into 2 different training groups that trained 2x$wk21 for 6 weeks. After the training intervention, average gluteus maximus muscle thickness increased by 9.4% in the deep squat group and 3.7% in the hip thrust Figure 4. Length-tension relationship of the biceps brachii. Adapted from Koo et al. (30), with permission. Muscle Hypertrophy Response VOLUME 00 | NUMBER 00 | MONTH 20226 Copyright © National Strength and Conditioning Association. Unauthorized reproduction of this article is prohibited. group (p 5 0.001 between groups), thus suggesting that the gluteus max- imus responds well to training at longer muscle lengths. As previously men- tioned, the Kubo et al. (32) squat study found that deep squat training enhanced gluteus muscle volume by 6.7 6 3.5%, whereas half squat training only increased gluteus volume by 2.26 2.6%. Moreover, Nakamura et al. (45) investigated subjects performing fly- wheel squats to 90 degrees of knee flexion 2x/week for 5 weeks and found no significant changes (p . 0.05) in gluteus maximus muscle thickness, whereas Yasuda et al. (76) found that full ROM leg press (90-08 knee flexion and 125-708 hip flexion) training for 12 weeks resulted in a 4.4% increase in gluteus muscle thickness. Although the length-tension relation- ship of the gluteus maximus is currently unknown, the above findings from Bar- balho et al. (3), Kubo et al. (32), Naka- mura et al. (45), and Yasuda et al. (76) would suggest that the gluteus is likely active on the descending portion of the length-tension curve and that a greater hip flexion angle will likely induce greater hypertrophic adaptations than a shallow hip flexion angle. In addition, Contreras et al. (11) discovered that the gluteus exhibits maximal activation in the top region of the barbell hip thrust exercise which suggests that the gluteus likely is not active on the ascending portion of the curve. Muscles that are active on the ascending portion of the curve often exhibit a trait known as “active insufficiency” in which they shorten to a degree in which little force production is taking place (60). This is exhibited in studies in which the gas- trocnemius, a biarticulate muscle well- known to experience active insuffi- ciency, displays a reduction in muscle activation during ankle plantar flexion with a flexed knee (23). Worthy of mention, the results of the Barbalho et al. (3) study are currently under scrutiny in a white paper sub- mitted by Vigotsky et al. (73) to the journal, SportRxiv. Although the article by Barbalho et al. (3) faces potential retraction, its results do not disagree with the findings from Kubo et al. (32) in which the gluteus maximus dis- played significantly greater hypertro- phic adaptations to training with longer muscle lengths. Future research is warranted to truly uncover the length-tension relationship of the glu- teus maximus because these findings are of great interest to sport athletes and physique athletes alike. BICEPS A recent project from Sato et al. (59) slightly mirrored that of Pinto et al. (55) in which elbow flexor muscle thick- ness was assessed before and after 2 unique biceps training programs. Sato et al. (59) recruited 32 male and female university students and divided them into 3 groups: a control group, a long muscle length group (0–508 elbow flex- ion; full elbow extension was considered 08 flexion), and a short muscle length group (80–1308 elbow flexion). Both training groups performed preacher curls for 3 sets of 10 repetitions in a pro- gressive loading fashion specific to maximal isometric strength at the joint position trained by each group. Training sessions were performed twice per week for 5 weeks and outcome measures consisted of eccentric, concentric, and isometric torque as well as muscle thickness at 50, 60, and 70% of the humerus length between the lateral ep- icondyle and the acromion. Although Pinto et al. (55) found similar muscle hypertrophy adaptations between ranges of motion for the biceps, Sato et al. (59) reported that the long muscle length group made significantly greater average gains in muscle thickness than the short muscle length group (8.9 6 3.9 vs. 3.4 6 2.7%; p , 0.01). At first, these results may seem at odds with the findings of Pinto et al. (55). However, subjects in the short muscle length group designed by Sato et al. (59) performed biceps curls from 80 to 1308 of elbow flexion, whereas the subjects in the Pinto et al. (55) study used an elbow flexion range of 50–1008. Therefore, it is plau- sible that training the biceps in the shortened range seen in the study by Sato et al. (59) may result in active insufficiency in the biceps and impaired hypertrophic adaptations, whereas the 50–1008 range used by Pinto et al. (55) may not. Certainly, more evidence is needed to verify this theory. A related, but unique, study from Nunes et al. (49) examined the effect of maximal load placement with respect to muscle length, rather than altering the range of motion in an exer- cise. The authors examined 35 healthy adults aged 23.7 6 5.3 years perform- ing either free weight preacher curl or cable preacher curl training. Both groups performed full range of motion training; however, the free weight training group, because of the inherent biomechanics of the exercise, would achieve peak tension at longer muscle lengths, whereas the cable training group would attain the highest tension with the muscle in a shortened state. After 10 weeks of thrice weekly train- ing, it was found that both groups experienced similar increases in biceps muscle thickness (7% for cable; 8% for free weights; p 5 0.346), thus strength- ening the theory that the biceps need no emphasis on training at longer mus- cle lengths to maximize hypertrophy. Another curious study from de Vas- concelos Costa et al. (14) compared 2 groups of untrained men performing either a nonvaried or a varied exercise program 3x$wk21 for 9 weeks. The researchers then investigated changes in MTat the proximal, medial, and dis- tal sites of the major muscle groups trained throughout the intervention. A key difference between groups about biceps training was that the nonvaried group solely performed traditional bar- bell curls, whereas the varied exercise group performed barbell curls, preacher curls, and incline dumbbell curls, which could certainly increase the trained muscle length of the biceps because of slight shoulder extension during the incline curl. Both groups exhibited significant changes in biceps MT from pre to post for both the medial (2.2 vs. 3.1%; nonvaried vs. var- ied, respectively) and distal sites (3.5 vs. 3.7%); however, only the varied exer- cise group displayed a significant Strength and Conditioning Journal | www.nsca-scj.com 7 Copyright © National Strength and Conditioning Association. Unauthorized reproduction of this article is prohibited. increase in proximal biceps MT (2.1 vs. 4.5%). Intriguingly, there were no sig- nificant between groups differences for any of the sites measured, even the proximal biceps (p 5 0.15).These results suggest that there may be sup- porting evidence for the inclusion of biceps exercises with greater shoulder extension angles; however, further research is needed. HAMSTRINGS Maeo et al. (37) investigated the effects of 12 weeks of seated or prone knee flexion training on 20 healthy adults aged 23.5 6 1.6 years. Participants performed seated leg flexions on one leg and prone leg flexions on the other. Both exercises involved a knee range of motion from 0 to 908; however, the seated leg curl also included a position of 908 of hip flexion, whereas the prone leg flexion condition involved 308 of hip flexion. After the intervention, it was found that the leg performing seated knee flexion training experienced a greater increase in muscle volume in 3 of the 4 hamstrings muscles than the group performing prone knee flexion training (14 vs. 9%; p # 0.010). Intriguingly, both groups exhibited a similar increase in the monoarticulate biceps femoris short head muscle (10 vs. 9%; p 5 0.190). Because 3 of the 4 heads of the hamstrings are biarticulate (semite- ndinosus, semimembranosus, and biceps femoris long head), these mus- cles are active on the plateau and de- scending regions of the length-tension curve (37,75), and thus, they likely respond greater to training at long muscle lengths. However, the biceps femoris short head is a monoarticulate Figure 5. Length-tension relationship of the latissimus dorsi. Adapted from Gerling and Brown (18), with permission. Figure 6. Length-tension relationship of the pectoralis. Adapted from Garner and Pandy (17), with permission. Muscle Hypertrophy Response VOLUME 00 | NUMBER 00 | MONTH 20228 Copyright © National Strength and Conditioning Association. Unauthorized reproduction of this article is prohibited. muscle because it only crosses the knee joint. Therefore, although the biceps femoris short head is also active on the plateau and descending regions of the length-tension curve (37,75), hip flexion (as occurs in seated knee flexion training) would have little influence on the length change in this muscle. OTHER MUSCLE GROUPS Although a small number of previous studies have endeavored to explore the unique adaptations to altering the range of motion used in strength train- ing exercises, not every major muscle group has been examined. Thus, this section serves to create a theoretical basis for application and provide routes for future research. Previous research has uncovered that the latissimus dorsi is active on both the plateau and descending regions of the length-tension curve (18) which would suggest that the latissimus dorsi is best trained at long muscle lengths; however, further research is needed to verify this theory. The length-tension relationship of the latissimus dorsi is shown in Figure 5. Previous research from Garner and Pandy (17) indicates that the pectoralis major is active on all portions of the length-tension curve. Thus, it is likely optimal to train this muscle group at longer muscle lengths because the pecto- rals may experience active insufficiency at shorter muscle lengths; however, these theories require further validation in research settings. The length-tension relationship of the pectoralis is shown in Figure 6. It is known that the deltoid muscle group contains 3 unique muscle heads, and intriguingly, the posterior head displays a unique length-tension curve compared with the other 2 heads of the deltoid. Indeed, research from Gar- ner and Pandy (17) suggests that all 3 heads of the deltoid muscle are active in the ascending and plateau region of Figure 7. Length-tension relationship of the deltoid major. Adapted from Garner and Pandy (17), with permission. Strength and Conditioning Journal | www.nsca-scj.com 9 Copyright © National Strength and Conditioning Association. Unauthorized reproduction of this article is prohibited. the curve; however, only the anterior and lateral deltoid reach the descend- ing portion of the curve. Therefore, it is likely that the posterior head of the deltoid need not be trained at longer muscle lengths for optimal hypertro- phy because its length-tension curve is similar to that of the biceps brachii (30). Similar to nearly every other mus- cle group, further research is needed to verify this hypothesis. The length- tension relationship of the deltoids is shown in Figure 7. Research has uncovered that all regions of the trapezius muscle are active on the ascending and plateau regions of the length-tension curve but not the descending portion (17). Although purely theoretical, it is certainly plausible that the trapezius group need not be trained at longer muscle lengths to achieve optimal hypertrophic adapta- tions, but this theory remains undis- turbed in the research field. The length-tension relationship for the trape- zius major is shown in Figure 8. The final major muscle group that is of interest to both fitness enthusiasts and athletes alike is the triceps surae. The triceps surae consist of the biarticulate gastrocnemius and monoarticulate soleus muscles. Owing to their inherent differences in anatomy, each muscle dis- plays a unique length-tension curve. Research from Chen and Delp (8) found that the soleus is only active on the pla- teau and descending regions of the length-tension curve. Conversely, previ- ous studies have reported that the gas- trocnemius is only active on the ascending and plateau portions of the curve (25). The position of the gastroc- nemius on the length-tension curve likely explains its active insufficiency during combined knee flexion and ankle plantar flexion movements (23). These data would suggest that the soleus muscle responds more favorably to using larger ranges of motion, whereas the gastroc- nemius should be primarily trained through heel raise variations without Figure 8. Length-tension relationship of the trapezius major. Adapted from Garner and Pandy (17), with permission. Muscle Hypertrophy Response VOLUME 00 | NUMBER 00 | MONTH 202210 Copyright © National Strength and Conditioning Association. Unauthorized reproduction of this article is prohibited. simultaneous knee flexion. The length- tension relationships of the soleus and gastrocnemius are shown in Figure 9. PRACTICAL APPLICATIONS Although research investigating the long-term hypertrophic effects of using different ranges of motion in strength training exercises is limited, a clear trend is certainly appearing that matches a muscle group’s long-term adaptations with its length-tension curve with respect to the ranges of motion used to train it. Indeed, neither the biceps (55) nor triceps (19,68) gar- nered additional benefits from using greater ranges of motion in strength training. Given that neither muscle group is active on the descending region of the length-tension curve, it is plausible that the range of motion used for training these muscles is not as important a training variable as other overload strategies. Conversely, research investigating the quadriceps (5,38) and hamstrings (37) shows that both muscle groups display a more sig- nificant hypertrophy response to train- ing with larger ranges of motion. Because both muscle groups are active on the descending portion of the length-tension curve, these muscles likely experience greater tension at long lengths, thus increasing the hyper- trophic stimulus from training with larger ranges of motion. This connection between concepts is certainly preliminary and requires fur- ther research; however, it provides support for using a muscle’s known length-tension properties and its rela- tionship with joint ranges of motion to design resistance training programs for optimizing muscular hypertrophy. Cer- tainly, the relationship between joint angle and muscle length needs further validation (and is influenced by a vari- ety of factors), but the strength and conditioning professional has limited resources for pursuing an applicable method of measuring muscle lengths during strength training exercises.Therefore, it is plausible that practi- tioners may prescribe greater joint ranges of motion to induce longer mus- cle lengths and optimally develop mus- cle groups that are active on the descending portion of the length- tension curve. Unfortunately, the above studies are the only inquiries into range of motion and long-term hypertrophic adaptations in resistance training. The theoretical connection between con- cepts presented in this article is Figure 9. Length-tension relationships of the soleus and gastrocnemius. Adapted from Chen and Delp (8) and Hoffman et al. (25), respectively. Strength and Conditioning Journal | www.nsca-scj.com 11 Copyright © National Strength and Conditioning Association. Unauthorized reproduction of this article is prohibited. undoubtedly preliminary and requires further research. However, it provides support for using a muscle’s known length-tension properties to design resis- tance training programs for optimizing muscular hypertrophy. Unfortunately, minimal studies have been performed investigating the effects of range of motion on muscle hypertrophy out- comes.Moreover, these studies have been performed on different muscle groups with unique length-tension curves; thus, muddying the oft-accepted theory that greater range of motion will result in superior muscle hypertrophy relative to smaller ranges of motion. Despite these setbacks, research into the length-tension relationship of nearly every major muscle group has been performed and Table 1 summarizes these findings. CONCLUSION Although preliminary, it is conceiv- able that applying a given muscle’s length-tension relationship to the joint ranges of motion used in resis- tance training programs may opti- mize hypertrophic adaptations. At the very least, this strategy could improve exercise selection for a given muscle group and reduce time spent on emphasizing ranges of motion that may be inconsequential to long-term outcomes. This senti- ment is underscored by research from Baroni et al. (4) in which full Table 1 Summary of L-T relationship for major muscle groups Muscle group Ascending Y/N Plateau Y/N Descending Y/N Applications References Gastrocnemius Yes Yes No May not necessitate training at longer lengths. Experiences active insufficiency at short lengths. (23,25) Soleus No Yes Yes Likely experiences greater hypertrophy at longer muscle lengths. (8) Quadriceps No Yes Yes Research has demonstrated greater hypertrophy at greater lengths. Cutoff may be at 908 knee flexion. (5,32,38,53,67) Hamstrings No Yes Yes All hamstrings muscles likely experience greater hypertrophy at longer muscle lengths. (37,75) Gluteus maximus No* Yes* Yes* Research has demonstrated that the gluteus maximus responds more favorably to large ROM training. True length-tension relationship unknown. (3,32,45,76) Latissimus dorsi No Yes Yes Likely experiences greater hypertrophy at longer muscle lengths. (18) Pectoralis major Yes Yes Yes Likely experiences greater hypertrophy at longer muscle lengths. May also experience active insufficiency. (17) Deltoid group Yes Yes Yes/No All 3 heads likely experience greater hypertrophy at longer lengths but may also experience active insufficiency. Posterior deltoid is not active on descending portion. (17) Trapezius Yes Yes No May not necessitate training at longer lengths. May experience active insufficiency at short lengths. (17) Biceps Yes Yes No May not necessitate training at longer lengths. Likely experiences active insufficiency at short lengths. (14,30,49,55,69) Triceps No Yes No Muscle length likely irrelevant for maximizing triceps hypertrophy. (19,44,68) Muscle Hypertrophy Response VOLUME 00 | NUMBER 00 | MONTH 202212 Copyright © National Strength and Conditioning Association. Unauthorized reproduction of this article is prohibited. range of motion elbow flexion train- ing was found to induce more muscle damage in the biceps than partial range of motion training. Muscle damage and the resultant soreness can negatively affect recovery and future training session quality, but the preliminary theory presented here suggests that there is no hyper- trophic benefit to using a full range of motion when training the biceps. To the strength and conditioning profes- sional limited by time or scheduling constraints, using these potential findings may offer useful insight that aids in program design. Importantly, the practitioner may also need to consider the inherent alter- ations in muscle and joint moments when modifying the range of motion used in resistance training exercises. It is generally accepted that for most muscles, increasing the flexion angle of a joint will result in a greater inter- nal moment, and thus, larger levels of tension experienced by the target muscle. Indeed, preliminary research would suggest that increasing the range of motion (and resultant inter- nal moment) enhances hypertrophic adaptations in some muscles but not others, which may embolden the practical usefulness of the length- tension curve in this aspect. Although the concept of joint and muscle moments likely interacts with the pre- sented theories relative to the length- tension curve, further research is war- ranted to ascertain the independence of each stimulus about hypertrophic outcomes. Relative to the other hard sciences, exercise and strength training research is a relatively new endeavor (20). New findings and theories are published virtually every day, often- times with inconclusive results that warrant further research. Unique training methods and theories are occasionally met with resistance from practitioners, especially because of the common use of untrained sub- jects in strength training literature. It is well-established that adaptations to training differ greatly between trained and untrained individuals (1,2) which can reduce the applicability of a study’s results. Furthermore, contradictory findings, such as seen between Bloom- quist et al. (5) and Pinto et al. (55), can add to the general confusion of the field when appropriate connections between muscle properties and their influence on adaptations are not made. It is difficult to identify holes in the research when it comes to strength training ranges of motion, associated muscle lengths, and long-term adapta- tions because the topic itself is a hole in the research and requires further investigation. Future research should endeavor to perform varying range of motion training with all the major muscle groups to assess the veracity of the theory presented herein. In addi- tion, using trained subjects in these investigations offer more valid insights for the practicing strength and condi- tioning professional. Finally, investiga- tions such as Stasinaki et al. (68) may be considered a gold standard in this realm in which ultrasonography was used to measure muscle fascicle length which validated the joint range of motion used in the training proto- col. Until the appropriate literature has been established, the theory pre- sented in this review may serve as the proverbial bridge between science and application for the strength and conditioning practitioner and fitness enthusiast alike. Conflicts of Interest and Source of Funding: The authors report no conflicts of interest and no source of funding. Charlie R. Ottinger is the Director of Human Perfor- mance at the Applied Science and Performance Institute. Matthew H. Sharp is the Vice President of Research at the Applied Science and Performance Institute. Matthew W. Stefan is a Research Scien- tist at the Applied Science and Performance Institute. Raad H. Gheith is a Research Scientist at the Applied Science and Performance Institute. Fernando de la Espriella is the Director of Crea- tives at the Applied Science and Performance Institute. Jacob M. Wilson is the Chief Executive Officer at the Applied Science and Performance Institute. 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