Bite force is critical to feeding success, especially in animals that crush strong, brittle foods. Maximum bite force is typically measured as one value per individual, but the force -length relationship of skeletal muscle suggests that each individual should possess a range of gape height-specific, and, therefore, prey size-specific, bite forces. We characterized the influence of prey size on pharyngeal jaw bite force in the snail-eating black carp (Mylopharyngodon piceus, family Cyprinidae), using feeding trials on artificial prey that varied independently in size and strength. We then measured jawclosing muscle lengths in vivo for each prey size, and then determined the force-length relationship of the same muscle in situ using tetanic stimulations. Maximum bite force was surprisingly high: the largest individual produced nearly 700 N at optimal muscle length. Bite force decreased on large and small prey, which elicited long and short muscle lengths, respectively, demonstrating that the force-length relationship of skeletal muscle results in prey size-specific bite force.
SUMMARYSand lances, fishes in the genus Ammodytes, exhibit a peculiar burrowing behavior in which they appear to swim rapidly into the substrate. They use posteriorly propagated undulations of the body to move in both water, a Newtonian fluid, and in sand, a nonNewtonian, granular substrate. In typical aquatic limbless locomotion, undulations of the body push against water, which flows because it is incapable of supporting the static stresses exerted by the animal, thus the undulations move in world space (slipping wave locomotion). In typical terrestrial limbless locomotion, these undulations push against substrate irregularities and move relatively little in world space (non-slipping wave locomotion). We used standard and X-ray video to determine the roles of slipping wave and non-slipping wave locomotion during burrowing in sand lances. We find that sand lances in water use slipping wave locomotion, similar to most aquatic undulators, but switch to non-slipping waves once they burrow. We identify a progression of three stages in the burrowing process: first, aquatic undulations similar to typical anguilliform locomotion (but without head yaw) push the head into the sand; second, more pronounced undulations of the aquatic portion of the body push most of the animal below ground; third, the remaining above-ground portion of the body ceases undulation and the subterranean portion takes over, transitioning to non-slipping wave locomotion. We find no evidence that sand lances use their body motions to fluidize the sand. Instead, as soon as enough of the body is underground, they undergo a kinematic shift and locomote like terrestrial limbless vertebrates.
The tradeoff between force and velocity in skeletal muscle is a fundamental constraint on vertebrate musculoskeletal design (form:function relationships). Understanding how and why different lineages address this biomechanical problem is an important goal of vertebrate musculoskeletal functional morphology. Our ability to answer questions about the different solutions to this tradeoff has been significantly improved by recent advances in techniques for quantifying musculoskeletal morphology and movement. Herein, we have three objectives: (1) review the morphological and physiological parameters that affect muscle function and how these parameters interact; (2) discuss the necessity of integrating morphological and physiological lines of evidence to understand muscle function and the new, high resolution imaging technologies that do so; and (3) present a method that integrates high spatiotemporal resolution motion capture (XROMM, including its corollary fluoromicrometry), high resolution soft tissue imaging (diceCT), and electromyography to study musculoskeletal dynamics in vivo. The method is demonstrated using a case study of in vivo primate hyolingual biomechanics during chewing and swallowing. A sensitivity analysis demonstrates that small deviations in reconstructed hyoid muscle attachment site location introduce an average error of 13.2% to in vivo muscle kinematics. The observed hyoid and muscle kinematics suggest that hyoid elevation is produced by multiple muscles and that fascicle rotation and tendon strain decouple fascicle strain from hyoid movement and whole muscle length. Lastly, we highlight current limitations of these techniques, some of which will likely soon be overcome through methodological improvements, and some of which are inherent. Anat Rec, 301:378-406, 2018. © 2018 Wiley Periodicals, Inc.
Aponeuroses are sheet-like elastic tendon structures that cover a portion of the muscle belly and act as insertion sites for muscle fibers while free tendons connect muscles to bones. During shortening contractions, free tendons are loaded in tension and lengthen due to the force acting longitudinally along the muscle’s line of action. In contrast, aponeuroses increase in length and width, suggesting that aponeuroses are loaded in directions along and orthogonal to the muscle’s line of action. Because muscle fibers are isovolumetric, they must expand radially as they shorten, potentially generating a force that increases aponeurosis width. We hypothesized that increases in aponeurosis width result from radial expansion of shortening muscle fibers. We tested this hypothesis by combining in situ muscle-tendon measurements with high-speed biplanar fluoroscopy measurements of the turkey’s lateral gastrocnemius (n = 6) at varying levels of isotonic muscle contractions. The change in aponeurosis width during periods of constant force depended on both the amount of muscle shortening and the magnitude of force production. At low to intermediate forces, aponeurosis width increased in direct proportion to fiber shortening. At high forces, aponeurosis width increased to a lesser extent or in some cases, decreased slightly during fiber shortening. Our results demonstrate that forces generated from radial expansion of shortening muscle fibers tend to drive increases in aponeurosis width, whereas longitudinal forces tend to decrease aponeurosis width. Ultimately, it is these two opposing forces that drive changes in aponeurosis width and alter series elastic stiffness during a muscle contraction.
SUMMARYPremaxillary protrusion in cypriniform fishes involves rotation of the kinethmoid, an unpaired skeletal element in the dorsal midline of the rostrum. No muscles insert directly onto the kinethmoid, so its rotation must be caused by the movement of other bones. In turn, the kinethmoid is thought to push on the ascending processes of the premaxillae, effecting protrusion. To determine the causes and effects of kinethmoid motion, we used XROMM (x-ray reconstruction of moving morphology) to measure the kinematics of cranial bones in common carp, Cyprinus carpio. Mean kinethmoid rotation was 83deg during premaxillary protrusion (18 events in 3 individuals). The kinethmoid rotates in a coordinated way with ventral translation of the maxillary bridge, and this ventral translation is likely driven primarily by the A1b muscle. Analyses of flexibility (variability between behaviors) and coordination (correlation between bones within a behavior) indicate that motion of the maxillary bridge, not the lower jaw, drives premaxillary protrusion. Thus, upper jaw protrusion is decoupled from lower jaw depression, allowing for two separate modes of protrusion, open mouth and closed mouth. These behaviors serve different functions: to procure food and to sort food, respectively. Variation in starting posture of the maxilla alone dictates which type of protrusion is performed; downstream motions are invariant. For closed mouth protrusion, a ventrally displaced maxillary starting posture causes kinethmoid rotation to produce more ventrally directed premaxillary protrusion. This flexibility, bestowed by the kinethmoid-maxillary bridge-A1b mechanism, one of several evolutionary novelties in the cypriniform feeding mechanism, may have contributed to the impressive trophic diversity that characterizes this speciose lineage.
The cellulose-rich walls that protect plant cells are difficult to digest, and therefore mechanical food processing is a key aspect of herbivory across vertebrates. Cell walls are typically broken down by translation of flattened teeth in the occlusal plane (i.e. grinding) as part of a complex, rhythmic chewing stroke. The grass carp, Ctenopharyngodon idella, is a voracious, invasive herbivorous fish that relies solely on its pharyngeal teeth, located in the back of the throat, for mechanical processing of plant material. Here, we describe the musculoskeletal anatomy of the pharyngeal jaws of grass carp and use XROMM to quantify chewing kinematics and muscle strain. The pharyngeal jaws are suspended in a sling of 11 muscles and maintain no bony articulation with any other skeletal elements in the head. The jaws bear long, serrated teeth that are worn during use into flattened tooth cusps. Our kinematic data show that this wear is the result of the teeth being elevated into occlusion against the basioccipital process and keratinous chewing pad, not tooth-on-tooth occlusion. Pharyngeal jaw elevation results from large strains in the jaw elevator muscle, the levator arcus branchialis V, to drive a pulleylike mechanism that rotates the jaws about a pivot point at the symphysis between the left and right pharyngeal jaws. These complex, rhythmic jaw rotations translate the teeth laterally across the chewing surface throughout the occlusion phase. The grass carp chewing system is strikingly similar in gross morphology and masticatory function to herbivorous chewing strategies in other vertebrates.
Generally behavioral neuroscience studies of the common marmoset employ adaptations of well-established training methods used with macaque monkeys. However, in many cases these approaches do not readily generalize to marmosets indicating a need for alternatives. Here we present the development of one such alternate: a platform for semiautomated, voluntary in-home cage behavioral training that allows for the study of naturalistic behaviors. We describe the design and production of a modular behavioral training apparatus using CAD software and digital fabrication. We demonstrate that this apparatus permits voluntary behavioral training and data collection throughout the marmoset’s waking hours with little experimenter intervention. Furthermore, we demonstrate the use of this apparatus to reconstruct the kinematics of the marmoset’s upper limb movement during natural foraging behavior. NEW & NOTEWORTHY The study of marmosets in neuroscience has grown rapidly and presents unique challenges. We address those challenges with an innovative platform for semiautomated, voluntary training that allows marmosets to train throughout their waking hours with minimal experimenter intervention. We describe the use of this platform to capture upper limb kinematics during foraging and to expand the opportunities for behavioral training beyond the limits of traditional training sessions. This flexible platform can easily incorporate other tasks.
Numerous therapeutics demonstrate optimal absorption or activity at specific sites in the gastrointestinal (GI) tract. Yet, safe, effective pill retention within a desired region of the GI remains an elusive goal. We report a safe, effective method for localizing magnetic pills. To ensure safety and efficacy, we monitor and regulate attractive forces between a magnetic pill and an external magnet, while visualizing internal dose motion in real time using biplanar videofluoroscopy. Real-time monitoring yields direct visual confirmation of localization completely noninvasively, providing a platform for investigating the therapeutic benefits imparted by localized oral delivery of new and existing drugs. Additionally, we report the in vitro measurements and calculations that enabled prediction of successful magnetic localization in the rat small intestines for 12 h. The designed system for predicting and achieving successful magnetic localization can readily be applied to any area of the GI tract within any species, including humans. The described system represents a significant step forward in the ability to localize magnetic pills safely and effectively anywhere within the GI tract. What our magnetic pill localization strategy adds to the state of the art, if used as an oral drug delivery system, is the ability to monitor the force exerted by the pill on the tissue and to locate the magnetic pill within the test subject all in real time. This advance ensures both safety and efficacy of magnetic localization during the potential oral administration of any magnetic pill-based delivery system. magnetic pills | retention | localization | drug delivery F or many orally administered pharmaceuticals, increased residence time in a particular region of the gastrointestinal (GI) tract would greatly improve their therapeutic benefit (1-3). Typically, physiological digestive processes govern the GI residence of standard pills. We have developed a magnet-based delivery system visualized by biplanar videofluoroscopy in vivo that yields real-time monitoring and control over the duration of GI residence of model magnetic pills in the small intestines of rats both as a tool for investigating the GI site-specific absorption and action of therapeutics and as a critical step towards enabling clinical localization of magnetic pills. Our system can safely and reliably retain drugs for a user-defined duration in any region of the GI with the ability to monitor and control the force applied by the orally ingested magnet to the intestinal wall.
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