Ontogenetic changes in bite force and gape in tufted capuchins.

Bite force and gape are two important performance metrics of the feeding system, and these metrics are inversely related for a given muscle size because of fundamental constraints in sarcomere length-tension relationships. How these competing performance metrics change in developing primates is largely unknown. Here, we quantified in vivo bite forces and gapes across ontogeny and examined these data in relation to body mass and cranial measurements in captive tufted capuchins, Sapajus spp. Bite force and gape were also compared across geometric and mechanical properties of mechanically challenging foods to investigate relationships between bite force, gape and food accessibility (defined here as the ability to breach shelled nuts). Bite forces at a range of gapes and feeding behavioral data were collected from a cross-sectional ontogenetic series of 20 captive and semi-wild tufted capuchins at the Núcleo de Procriação de Macacos-Prego Research Center in Araçatuba, Brazil. These data were paired with body mass, photogrammetric measures of jaw length and facial width, and food geometric and material properties. Tufted capuchins with larger body masses had absolutely higher in vivo bite forces and gapes, and animals with wider faces had absolutely higher bite forces. Bite forces and gapes were significantly smaller in juveniles compared with subadults and adults. These are the first primate data to empirically demonstrate the gapes at which maximum active bite force is generated and to demonstrate relationships to food accessibility. These data advance our understanding of how primates meet the changing performance demands of the feeding system during development.

Fiber-type phenotype of the jaw-closing muscles in Gorilla gorilla, Pan troglodytes, and Pan paniscus: A test of the Frequent Recruitment Hypothesis.

Skeletal muscle fiber types are important determinants of the contractile properties of muscle fibers, such as fatigue resistance and shortening velocity. Yet little is known about how jaw-adductor fiber types correlate with feeding behavior in primates. Compared with chimpanzees and bonobos, gorillas spend a greater percentage of their daily time feeding and shift to herbaceous vegetation when fruits are scarce. We thus used the African apes to test the hypothesis that chewing with unusually high frequency is correlated with the expression in the jaw adductors of a high proportion of type 1 (slow, fatigue-resistant) fibers at the expense of other fiber types (the Frequent Recruitment Hypothesis). We used immunohistochemistry to determine the presence and distribution of the four major myosin heavy chain (MHC) isoforms in the anterior superficial masseter (ASM), superficial anterior temporalis, and deep anterior temporalis of four Gorilla gorilla, two Pan paniscus, and four Pan troglodytes. Serial sections were stained against slow (MHC-1/-α-cardiac) and fast (MHC-2/-M) fibers. Fibers were counted and scored for staining intensity, and fiber cross-sectional areas (CSAs) were measured and used to estimate percentage of CSA of each MHC isoform. Hybrid fibers accounted for nearly 100% of fiber types in the masseter and temporalis of all three species, resulting in three main hybrid phenotypes. As predicted, the gorilla ASM and deep anterior temporalis comprised a greater percentage of CSA of the slower, fatigue-resistant hybrid fiber type, significantly so for the ASM (p = 0.015). Finally, the results suggest that fiber phenotype of the chewing muscles contributes to behavioral flexibility in ways that would go undetected in paleontological studies relying solely on morphology of the bony masticatory apparatus.

Biomechanics of the mandible of Macaca mulatta during the power stroke of mastication: Loading, deformation, and strain regimes and the impact of food type.

Mandible morphology has yet to yield definitive information on primate diet, probably because of poor understanding of mandibular loading and strain regimes, and overreliance on simple beam models of mandibular mechanics. We used a finite element model of a macaque mandible to test hypotheses about mandibular loading and strain regimes and relate variation in muscle activity during chewing on different foods to variation in strain regimes. The balancing-side corpus is loaded primarily by sagittal shear forces and sagittal bending moments. On the working side, sagittal bending moments, anteroposterior twisting moments, and lateral transverse bending moments all reach similar maxima below the bite point; sagittal shear is the dominant loading regime behind the bite point; and the corpus is twisted such that the mandibular base is inverted. In the symphyseal region, the predominant loading regimes are lateral transverse bending and negative twisting about a mediolateral axis. Compared with grape and dried fruit chewing, nut chewing is associated with larger sagittal and transverse bending moments acting on balancing- and working-side mandibles, larger sagittal shear on the working side, and larger twisting moments about vertical and transverse axes in the symphyseal region. Nut chewing is also associated with higher minimum principal strain magnitudes in the balancing-side posterior ramus; higher sagittal shear strain magnitudes in the working-side buccal alveolar process and the balancing-side oblique line, recessus mandibulae, and endocondylar ridge; and higher transverse shear strains in the symphyseal region, the balancing-side medial prominence, and the balancing-side endocondylar ridge. The largest food-related differences in maximum principal and transverse shear strain magnitudes are in the transverse tori and in the balancing-side medial prominence, extramolar sulcus, oblique line, and endocondylar ridge. Food effects on the strain regime are most salient in areas not traditionally investigated, suggesting that studies seeking dietary effects on mandible morphology might be looking in the wrong places.

The dental microwear of hard-object feeding in laboratory Sapajus apella and its implications for dental microwear formation.

OBJECTIVES: This study seeks to determine if (a) consumption of hard food items or a mixture of food items leads to the formation of premolar or molar microwear in laboratory capuchin monkeys (Sapajus apella) in one feeding session and (b) rates of microwear formation are associated with the number of food items consumed. MATERIALS AND METHODS: Five adult male capuchins were used in two experiments, one where they were fed unshelled Brazil nuts, and the other where they were fed a mixture of food items. Dental impressions were taken before and after each feeding session. Epoxy casts made from those impressions then were used in SEM analyses of rates of microwear formation. Upper and lower premolars and molars were analyzed. Qualitative comparisons were made and Spearman's rank-order correlations used to examine the relationship between rates of microwear formation and number of Brazil nuts consumed. RESULTS: Premolars and molars generally showed new microwear in the form of pits and scratches. However, the incidence of those features was low (0-6%). Rates of microwear formation were highest during the consumption of Brazil nuts. DISCUSSION: Variations in the rate of microwear formation on the premolars likely reflected patterns of ingestion whereas consistency in the rate of microwear on the molars likely reflected patterns of chewing. While dental microwear formation seemed to be correlated with the number of hard objects consumed, rates did differ between individuals. Differences in results between the two experiments demonstrate some of the limitations in our knowledge of dental microwear formation.

Joint angular excursions during cyclical behaviors differ between tetrapod feeding and locomotor systems.

Tetrapod musculoskeletal diversity is usually studied separately in feeding and locomotor systems. However, comparisons between these systems promise important insight into how natural selection deploys the same basic musculoskeletal toolkit - connective tissues, bones, nerves and skeletal muscle - to meet the differing performance criteria of feeding and locomotion. In this study, we compare average joint angular excursions during cyclic behaviors - chewing, walking and running - in a phylogenetic context to explore differences in the optimality criteria of these two systems. Across 111 tetrapod species, average limb-joint angular excursions during cyclic locomotion are greater and more evolutionarily labile than those of the jaw joint during cyclic chewing. We argue that these findings reflect fundamental functional dichotomies between tetrapod locomotor and feeding systems. Tetrapod chewing systems are optimized for precise application of force over a narrower, more controlled and predictable range of displacements, the principal aim being to fracture the substrate, the size and mechanical properties of which are controlled at ingestion and further reduced and homogenized, respectively, by the chewing process. In contrast, tetrapod limbed locomotor systems are optimized for fast and energetically efficient application of force over a wider and less predictable range of displacements, the principal aim being to move the organism at varying speeds relative to a substrate whose geometry and mechanical properties need not become more homogeneous as locomotion proceeds. Hence, the evolution of tetrapod locomotor systems has been accompanied by an increasing diversity of limb-joint excursions, as tetrapods have expanded across a range of locomotor substrates and environments.

The Source and the Course of the Articular Branches to the T4-T8 Zygapophysial Joints.

OBJECTIVE: To define the source and the course of the articular branches to the midthoracic zygapophysial ("z") joints. DESIGN: Cadaveric dissection. SETTING: The Gross Anatomy Laboratory of the Duke University School of Medicine. SUBJECTS: Ten human cadaveric thoraces. METHODS: Gross and stereoscopic dissection of dorsal rami T4-T8 was performed bilaterally on 10 adult embalmed cadavers. The medial and lateral branches were traced to their origins from the dorsal rami, and the course of the articular nerves was documented through digital photography. Radio-opaque wire (20 gauge) was applied to the nerves. Fluoroscopic images were obtained to delineate their radiographic course with respect to osseous landmarks. RESULTS: Forty-eight inferior articular branches were identified. Three (6.3%) originated from the medial branch and 44 (91.7%) from the dorsal ramus. One was indeterminate. Fifty-one superior articular branches were identified. Eight (15.7%) originated from the medial branch and 43 (84.3%) from the dorsal ramus. In 12% of cases (6/50), there was side-to-side asymmetry in the origins of the articular branches. Nerves were commonly suspended in the intertransverse space. The articular branches contacted an osseous structure in only 39% of cases. As previously reported, a "descending branch" was not identified in any specimen. CONCLUSIONS: Articular branches to the T4-T8 z-joints have substantial inter- and intraspecimen variability of origin. They typically arise from the dorsal ramus rather than the medial branch and frequently do not contact any osseous structure to allow percutaneous needle placement.

Fiber type composition of epaxial muscles is geared toward facilitating rapid spinal extension in the leaper Galago senegalensis.

OBJECTIVES: We hypothesized that the vertical leaper Galago senegalensis will have epaxial extensor muscles with a fast fiber phenotype to facilitate rapid spinal extension during leaping in comparison to the slow-moving quadruped Nycticebus coucang. To test this, we determined the percentage of fiber cross-sectional area (%CSA) devoted to Type 2 fibers in epaxial muscles of G. senegalensis compared to those of N. coucang. MATERIALS AND METHODS: Immunohistochemistry was used to identify Type 1, Type 2, and hybrid fibers in iliocostalis, longissimus, and multifidus muscles of G. senegalensis (n = 3) and N. coucang (n = 3). Serial muscle sections were used to estimate and compare proportions, cross-sectional areas (CSAs), and %CSAs of Type 1, Type 2, and hybrid fibers between species. RESULTS: Epaxial muscles of G. senegalensis were comprised predominantly of Type 2 fibers with large CSAs (%CSA range ≈ 83-94%; range of mean CSA = 1,218-1,586 µm2 ). N. coucang epaxial muscles were comprised predominantly Type 1 fibers with large CSAs (%CSA range ≈ 69-77%; range of mean CSA = 983-1,220 µm2 ). DISCUSSION: The predominance of Type 2 fibers in G. senegalensis epaxial muscles facilitates rapid muscle excursion and spinal extension during leaping, and is consistent with their relatively long muscle fibers. The predominance of Type 1 fibers in N. coucang epaxial muscles may aid in maintaining stable postures during bridging and cantilevering behaviors characteristic of slow-climbing. These histochemical characteristics highlight the major divergent locomotor repertoires of G. senegalensis and N. coucang.

Participation, representation, and shared experiences of women scholars in biological anthropology.

American Association of Physical Anthropologists (AAPA) membership surveys from 1996 and 1998 revealed significant gender disparities in academic status. A 2014 follow-up survey showed that gender equality had improved, particularly with respect to the number of women in tenure-stream positions. However, although women comprised 70% of AAPA membership at that time, the percentage of women full professors remained low. Here, we continue to consider the status of women in biological anthropology by examining the representation of women through a quantitative analysis of their participation in annual meetings of the AAPA during the past 20 years. We also review the programmatic goals of the AAPA Committee on Diversity Women's Initiative (COD-WIN) and provide survey results of women who participated in COD-WIN professional development workshops. Finally, we examine the diversity of women's career paths through the personal narratives of 14 women biological anthropologists spanning all ranks from graduate student to Professor Emeritus. We find that over the past 20 years, the percentage of women first authors of invited symposia talks has increased, particularly in the sub-disciplines of bioarchaeology, genetics, and paleoanthropology. The percentage of women first authors on contributed talks and posters has also increased. However, these observed increases are still lower than expected given the percentage of graduate student women and women at the rank of assistant and associate professor. The personal narratives highlight first-hand the impact of mentoring on career trajectory, the challenges of achieving work-life satisfaction, and resilience in the face of the unexpected. We end with some suggestions for how to continue to improve equality and equity for women in biological anthropology.

Scaling of rotational inertia of primate mandibles.

The relative importance of pendulum mechanics and muscle mechanics in chewing dynamics has implications for understanding the optimality criteria driving the evolution of primate feeding systems. The Spring Model (Ross et al., 2009b), which modeled the primate chewing system as a forced mass-spring system, predicted that chew cycle time would increase faster than was actually observed. We hypothesized that if mandibular momentum plays an important role in chewing dynamics, more accurate estimates of the rotational inertia of the mandible would improve the accuracy with which the Spring Model predicts the scaling of primate chew cycle period. However, if mass-related momentum effects are of negligible importance in the scaling of primate chew cycle period, this hypothesis would be falsified. We also predicted that greater "robusticity" of anthropoid mandibles compared with prosimians would be associated with higher moments of inertia. From computed tomography scans, we estimated the scaling of the moment of inertia (Ij) of the mandibles of thirty-one species of primates, including 22 anthropoid and nine prosimian species, separating Ij into the moment about a transverse axis through the center of mass (Ixx) and the moment of the center of mass about plausible axes of rotation. We found that across primates Ij increases with positive allometry relative to jaw length, primarily due to positive allometry of jaw mass and Ixx, and that anthropoid mandibles have greater rotational inertia compared with prosimian mandibles of similar length. Positive allometry of Ij of primate mandibles actually lowers the predictive ability of the Spring Model, suggesting that scaling of primate chew cycle period, and chewing dynamics in general, are more strongly influenced by factors other than scaling of inertial properties of the mandible, such as the dynamic properties of the jaw muscles and neural control. Differences in cycle period scaling between chewing and locomotion systems reinforce the suggestion that displacement and force control are more important in the design of feeding systems than energetics and speed.

In vivo bone strain in the mandibular corpus of Sapajus during a range of oral food processing behaviors.

It has been hypothesized that mandibular corpus morphology of primates is related to the material properties of the foods that they chew. However, chewing foods with different material properties is accompanied by low levels of variation in mandibular strain patterns in macaques. We hypothesized that if variation in primate mandible form reflects adaptations to feeding on foods with different material and geometric properties, then this variation will be driven primarily by differences in oral food processing behavior rather than differences in chewing per se. To test this hypothesis, we recorded in vivo bone strain data from the lateral and medial surfaces of the mandibular corpus during complete feeding sequences in three adult male Sapajus as they fed on foods with a range of sizes and material properties. We assessed whether variation in mandibular corpus strain regimes is associated with variation in feeding behaviors and/or chewing on different foods, and we quantified the relative variation in mandibular corpus strain regimes associated with chewing on foods of different material properties versus a range of oral food processing behaviors (incisor, premolar, and molar biting; pulling on incisors; mastication). Feeding behavior had a significant effect on mandibular corpus strain regimes, as did chewing side and the cycle number in a feeding sequence. However, food type had weaker effects and usually only through interaction effects with chewing side and/or cycle type. Strain regimes varied most across different chew sides, then across different behaviors, and lastly between mastication cycles on different foods. Strain magnitudes associated with premolar, molar, and incisor biting were larger than those recorded during mastication. These data suggest that intra- and inter-specific variation in mandible morphology is a trade-off between performance requirements of different oral food processing behaviors and of variation in chewing side, with direct effects of food type being less important.

Primate dietary ecology in the context of food mechanical properties.

Substantial variation exists in the mechanical properties of foods consumed by primate species. This variation is known to influence food selection and ingestion among non-human primates, yet no large-scale comparative study has examined the relationships between food mechanical properties and feeding strategies. Here, we present comparative data on the Young's modulus and fracture toughness of natural foods in the diets of 31 primate species. We use these data to examine the relationships between food mechanical properties and dietary quality, body mass, and feeding time. We also examine the relationship between food mechanical properties and categorical concepts of diet that are often used to infer food mechanical properties. We found that traditional dietary categories, such as folivory and frugivory, did not faithfully track food mechanical properties. Additionally, our estimate of dietary quality was not significantly correlated with either toughness or Young's modulus. We found a complex relationship among food mechanical properties, body mass, and feeding time, with a potential interaction between median toughness and body mass. The relationship between mean toughness and feeding time is straightforward: feeding time increases as toughness increases. However, when considering median toughness, the relationship with feeding time may depend upon body mass, such that smaller primates increase their feeding time in response to an increase in median dietary toughness, whereas larger primates may feed for shorter periods of time as toughness increases. Our results emphasize the need for additional studies quantifying the mechanical and chemical properties of primate diets so that they may be meaningfully compared to research on feeding behavior and jaw morphology.

Epaxial muscle fiber architecture favors enhanced excursion and power in the leaper Galago senegalensis.

Galago senegalensis is a habitual arboreal leaper that engages in rapid spinal extension during push-off. Large muscle excursions and high contraction velocities are important components of leaping, and experimental studies indicate that during leaping by G. senegalensis, peak power is facilitated by elastic storage of energy. To date, however, little is known about the functional relationship between epaxial muscle fiber architecture and locomotion in leaping primates. Here, fiber architecture of select epaxial muscles is compared between G. senegalensis (n = 4) and the slow arboreal quadruped, Nycticebus coucang (n = 4). The hypothesis is tested that G. senegalensis exhibits architectural features of the epaxial muscles that facilitate rapid and powerful spinal extension during the take-off phase of leaping. As predicted, G. senegalensis epaxial muscles have relatively longer, less pinnate fibers and higher ratios of tendon length-to-fiber length, indicating the capacity for generating relatively larger muscle excursions, higher whole-muscle contraction velocities, and a greater capacity for elastic energy storage. Thus, the relatively longer fibers and higher tendon length-to-fiber length ratios can be functionally linked to leaping performance in G. senegalensis. It is further predicted that G. senegalensis epaxial muscles have relatively smaller physiological cross-sectional areas (PCSAs) as a consequence of an architectural trade-off between fiber length (excursion) and PCSA (force). Contrary to this prediction, there are no species differences in relative PCSAs, but the smaller-bodied G. senegalensis trends towards relatively larger epaxial muscle mass. These findings suggest that relative increase in muscle mass in G. senegalensis is largely attributable to longer fibers. The relative increase in erector spinae muscle mass may facilitate sagittal flexibility during leaping. The similarity between species in relative PCSAs provides empirical support for previous work linking osteological features of the vertebral column in lorisids with axial stability and reduced muscular effort associated with slow, deliberate movements during anti-pronograde locomotion.

Jaw-muscle architecture and mandibular morphology influence relative maximum jaw gapes in the sexually dimorphic Macaca fascicularis.

Maximum jaw gape is a performance variable related to feeding and non-feeding oral behaviors, such as canine gape displays, and is influenced by several factors including jaw-muscle fiber architecture, muscle position on the skull, and jaw morphology. Maximum gape, jaw length, and canine height are strongly correlated across catarrhine primates, but relationships between gape and other aspects of masticatory apparatus morphology are less clear. We examine the effects of jaw-adductor fiber architecture, jaw-muscle leverage, and jaw form on gape in an intraspecific sample of sexually dimorphic crab-eating macaques (Macaca fascicularis). As M. fascicularis males have relatively larger maximum gapes than females, we predict that males will have muscle and jaw morphologies that facilitate large gape, but these morphologies may come at some expense to bite force. Male crab-eating macaques have relatively longer jaw-muscle fibers, masseters with decreased leverage, and temporomandibular joint morphologies that facilitate the production of wide gapes. Because relative canine height is correlated with maximum gape in catarrhines, and males have relatively longer canines than females, these results support the hypothesis that male M. fascicularis have experienced selection to increase maximum gape. The sexes do not differ in relative masseter physiologic cross-sectional area (PCSA), but males compensate for a potential trade-off between muscle excursion versus muscle force with increased temporalis weight and PCSA. This musculoskeletal configuration is likely functionally significant for behaviors involving aggressive canine biting and displays in male M. fascicularis and provides additional evidence supporting the multifactorial nature of the catarrhine masticatory apparatus. Our results have implications for the evolution of craniofacial morphology in catarrhine primates and reinforce the importance of evaluating additional factors other than feeding behavior and diet in analyses of masticatory apparatus form, function, and evolution.

Locomotor activity influences muscle architecture and bone growth but not muscle attachment site morphology.

The ability to make behavioural inferences from skeletal remains is critical to understanding the lifestyles and activities of past human populations and extinct animals. Muscle attachment site (enthesis) morphology has long been assumed to reflect muscle strength and activity during life, but little experimental evidence exists to directly link activity patterns with muscle development and the morphology of their attachments to the skeleton. We used a mouse model to experimentally test how the level and type of activity influences forelimb muscle architecture of spinodeltoideus, acromiodeltoideus, and superficial pectoralis, bone growth rate and gross morphology of their insertion sites. Over an 11-week period, we collected data on activity levels in one control group and two experimental activity groups (running, climbing) of female wild-type mice. Our results show that both activity type and level increased bone growth rates influenced muscle architecture, including differences in potential muscular excursion (fibre length) and potential force production (physiological cross-sectional area). However, despite significant influences on muscle architecture and bone development, activity had no observable effect on enthesis morphology. These results suggest that the gross morphology of entheses is less reliable than internal bone structure for making inferences about an individual's past behaviour.

Jaw-muscle force and excursion scale with negative allometry in platyrrhine primates.

OBJECTIVES: Platyrrhines span two orders of magnitude in body size and are characterized by diverse feeding behaviors and diets. While size plays an important role in primate feeding behavior and masticatory apparatus morphology, we know little about size-correlated changes in the force-generating (physiologic cross-sectional area; PCSA) and excursion/stretch (fiber length; Lf ) capabilities of the jaw-closing muscles in platyrrhines. METHODS: We examined scaling relationships of the superficial masseter and temporalis muscles in 21 platyrrhine species. Previous work suggests that larger platyrrhines are at a mechanical disadvantage for generating bite forces compared with smaller platyrrhines. We hypothesize that scaling of jaw-muscle fiber architecture counters this size-correlated decrease in mechanical advantage. Thus, we predicted that jaw-muscle PCSAs and muscle weights scale with positive allometry while Lf s scale with negative allometry, relative to load-arm estimates for incisor/molar biting and chewing. RESULTS: Jaw-muscle PCSAs and Lf s appear to scale with negative allometry relative to load-arm estimates and body size. Negative allometry of jaw-muscle weights partially accounts for the size-correlated decreases in PCSA and Lf . Estimates of bite force also scale with negative allometry. CONCLUSION: Large-bodied platyrrhines (e.g., Alouatta) are at a relative disadvantage for generating jaw-muscle and bite force as well as jaw-muscle stretch, compared with smaller species (e.g., Callithrix). The net effect is that larger platyrrhines likely produce relatively smaller maximal bite forces compared with smaller taxa. Relative to small- and intermediate-sized platyrrhines, large-bodied platyrrhines feed on some of the least mechanically challenging foods, consistent with the size-correlated decrease in relative muscle and bite forces across the clade. Am J Phys Anthropol, 2015. © 2015 Wiley Periodicals, Inc. Am J Phys Anthropol 158:242-256, 2015. © 2015 Wiley Periodicals, Inc.

Eccentric training for prevention of hamstring injuries may depend on intervention compliance: a systematic review and meta-analysis.

BACKGROUND: Hamstring injury is a prevalent muscle injury in sports. Inconclusive evidence exists for eccentric hamstring strengthening to prevent hamstring injuries. One reason for this discrepancy may be the influence intervention non-compliance has on individual study estimates, and therefore pooled estimates. OBJECTIVE: This systematic review aims to determine the effect of eccentric hamstring strengthening on the risk of hamstring injury and quantitatively explores the impact of intervention non-compliance on the precision, heterogeneity and strength of pooled estimates. METHODS: A computer-assisted literature search of Medline, CINAHL, Cochrane, EMBASE, AMED, SportDiscus and PEDro databases was conducted with keywords related to eccentric strengthening and hamstring injury. The search was conducted from the end of a previous comprehensive review forward (1 December 2008 to 31 December 2013). Random-effects models were used for both main effects and a sensitivity analysis. Pooled estimate precision was measured with a confidence limit ratio (confidence limit ratio (CLR); upper limit divided by the lower limit) and heterogeneity was assessed with I(2), Cochrane's-Q and τ(2). A protocol was not registered for this review. RESULTS: Four out of 349 studies met the inclusion criteria. In main effects analysis, eccentric hamstring training did not reduce the risk of hamstring injury (risk ratio [RR]=0.59 ((95% CI 0.24 to 1.44)). This estimate was imprecise (CLR=6.0) with significant heterogeneity (p value 0.02, 69.6% variation and t(2)=0.57). Subjects compliant with eccentric strengthening had a significant (RR=0.35 ((95% CI 0.23 to 0.55)) reduction in hamstring injuries. This estimate was precise (CLR=2.4) and homogenous (p value=0.38, 2.8% variation and t(2)=0.007). CONCLUSIONS: The null-biased effect in using intent-to-treat methods from intervention non-compliance has a substantial impact on the precision, heterogeneity and the direction and strength of pooled estimates. Eccentric strengthening, with good compliance, appears to be successful in prevention of hamstring injury.