Soft skeletons transmit force with variable gearing.

A hydrostatic skeleton allows a soft body to transmit muscular force via internal pressure. A human's tongue, an octopus' arm and a nematode's body illustrate the pervasive presence of hydrostatic skeletons among animals, which has inspired the design of soft engineered actuators. However, there is a need for a theoretical basis for understanding how hydrostatic skeletons apply mechanical work. We therefore modeled the shape change and mechanics of natural and engineered hydrostatic skeletons to determine their mechanical advantage (MA) and displacement advantage (DA). These models apply to a variety of biological structures, but we explicitly consider the tube feet of a sea star and the body segments of an earthworm, and contrast them with a hydraulic press and a McKibben actuator. A helical winding of stiff, elastic fibers around these soft actuators plays a critical role in their mechanics by maintaining a cylindrical shape, distributing forces throughout the structure and storing elastic energy. In contrast to a single-joint lever system, soft hydrostats exhibit variable gearing with changes in MA generated by deformation in the skeleton. We found that this gearing is affected by the transmission efficiency of mechanical work (MA×DA) or, equivalently, the ratio of output to input work. The transmission efficiency changes with the capacity to store elastic energy within helically wrapped fibers or associated musculature. This modeling offers a conceptual basis for understanding the relationship between the morphology of hydrostatic skeletons and their mechanical performance.

Synthetic analysis of trophic diversity and evolution in Enantiornithes with new insights from Bohaiornithidae.

Enantiornithines were the dominant birds of the Mesozoic, but understanding of their diet is still tenuous. We introduce new data on the enantiornithine family Bohaiornithidae, famous for their large size and powerfully built teeth and claws. In tandem with previously published data, we comment on the breadth of enantiornithine ecology and potential patterns in which it evolved. Body mass, jaw mechanical advantage, finite element analysis of the jaw, and traditional morphometrics of the claws and skull are compared between bohaiornithids and living birds. We find bohaiornithids to be more ecologically diverse than any other enantiornithine family: Bohaiornis and Parabohaiornis are similar to living plant-eating birds; Longusunguis resembles raptorial carnivores; Zhouornis is similar to both fruit-eating birds and generalist feeders; and Shenqiornis and Sulcavis plausibly ate fish, plants, or a mix of both. We predict the ancestral enantiornithine bird to have been a generalist which ate a wide variety of foods. However, more quantitative data from across the enantiornithine tree is needed to refine this prediction. By the Early Cretaceous, enantiornithine birds had diversified into a variety of ecological niches like crown birds after the K-Pg extinction, adding to the evidence that traits unique to crown birds cannot completely explain their ecological success.

The birds living in the world today are only a small part of the larger bird family tree. Around 120 to 65 million years ago, when dinosaurs and other large reptiles roamed the world, the ancestors of modern-day birds were actually rather rare. Instead, another now extinct group of birds called the Enantiornithes (meaning "opposite birds") were the most common birds. Many researchers believe that Enantiornithes may have filled similar roles in ancient ecosystems as living birds do today. For example, some may have hunted other birds or animals, while some may have eaten only plants. Some may have specialized at eating a few specific foods while others may have been 'generalists' that ate many different foods. However, some of the physical features of Enantiornithes set them apart from modern-day birds. For example, unlike living birds, Enantiornithes had teeth and their wings were also constructed very differently. Previous studies suggest that one group of these extinct birds most likely ate insects and another group most likely ate fish, but it remains unclear what variety of foods opposite birds as a whole may have consumed. Miller et al. compared the jaws, claws and various other physical features of fossils from six additional species of opposite birds with the skeletons of modern birds to infer what the diets of these opposite birds may have been. This approach revealed that Enantiornithes may have had a wide variety of different diets. The researchers found that two species probably ate plants, another species most likely ate meat, and another one likely ate a mixture of both. With a large sample across Enantiornithes, Miller et al. were able to predict the diet of their common ancestor. They found the common ancestor to most likely be a 'generalist' eating variety of foods and that some species subsequently evolved to have more specialist diets. Opposite birds probably played many different roles in ecosystems, like living birds do today. Therefore, a better understanding how Enantiornithes evolved may shed light on the factors that have influenced the evolution of modern-day birds. This may aid future conservation efforts to target birds whose descendants may be able to take up the ecological roles of other species that go extinct.

Diet of Mesozoic toothed birds (Longipterygidae) inferred from quantitative analysis of extant avian diet proxies.

BACKGROUND: Birds are key indicator species in extant ecosystems, and thus we would expect extinct birds to provide insights into the nature of ancient ecosystems. However, many aspects of extinct bird ecology, particularly their diet, remain obscure. One group of particular interest is the bizarre toothed and long-snouted longipterygid birds. Longipterygidae is the most well-understood family of enantiornithine birds, the dominant birds of the Cretaceous period. However, as with most Mesozoic birds, their diet remains entirely speculative. RESULTS: To improve our understanding of longipterygids, we investigated four proxies in extant birds to determine diagnostic traits for birds with a given diet: body mass, claw morphometrics, jaw mechanical advantage, and jaw strength via finite element analysis. Body mass of birds tended to correspond to the size of their main food source, with both carnivores and herbivores splitting into two subsets by mass: invertivores or vertivores for carnivores, and granivores + nectarivores or folivores + frugivores for herbivores. Using claw morphometrics, we successfully distinguished ground birds, non-raptorial perching birds, and raptorial birds from one another. We were unable to replicate past results isolating subtypes of raptorial behaviour. Mechanical advantage was able to distinguish herbivorous diets with particularly high values of functional indices, and so is useful for identifying these specific diets in fossil taxa, but overall did a poor job of reflecting diet. Finite element analysis effectively separated birds with hard and/or tough diets from those eating foods which are neither, though could not distinguish hard and tough diets from one another. We reconstructed each of these proxies in longipterygids as well, and after synthesising the four lines of evidence, we find all members of the family but Shengjingornis (whose diet remains inconclusive) most likely to be invertivores or generalist feeders, with raptorial behaviour likely in Longipteryx and Rapaxavis. CONCLUSIONS: This study provides a 20% increase in quantitatively supported fossil bird diets, triples the number of diets reconstructed in enantiornithine species, and serves as an important first step in quantitatively investigating the origins of the trophic diversity of living birds. These findings are consistent with past hypotheses that Mesozoic birds occupied low trophic levels.

Morphological, functional, and phylogenetic aspects of the head capsule of the cockroach Ergaula capucina (Insecta/Blattodea).

Background: Cockroaches are usually typical omnivorous detritivores and their cephalic morphology is considered to be ancestral in various aspects. Thus, several studies addressed the morphology and function of the blattodean head, and the cockroach usually serves as a model for standard mouthparts in text books. However, so far only two of the three major lineages of Blattodea have been studied and no detailed information for the head of any Corydioidea was available. The present study closes this gap by providing a detailed morphological description of the head of Ergaula capucina, studying some important functional parameters of the mandible and discussing it in a phylogenetic framework. Methods: The cephalic morphology of Ergaula studied in detail using a broad set of different techniques including digital microscopy, µ-computed tomography, and 3-dimensional reconstructions. Concerning the functional morphology of the mandible, we compared the volume and effective cross sections of the eight compartments of the primary mandibular adductor muscle for Ergaula, Blattella germanica, and Salganea rossi and measured the mechanical advantage, i.e., the force transmission ratio for all teeth of the mandible of Ergaula. Results: The head capsule of Ergaula is characterized by a strong sexual dimorphism and typical orthopteran mouthparts. It resembles the head capsule of other roaches in several respects and confirms oesotendons, the reduction of the mesal occelus, and bipartite M. verticopharyngealis and M. hypopharyngosalivaris as blattodean apomorphies. But it also shows some unique adaptations. It is the first described cockroach that lacks the dorsal tentorial arms which has various consequences for the cephalic musculature. On the maxillary lacinia, Ergaula is the first described blattodean to show strong and blunt setae instead of a lacinula, which might be homologues to the dentisetae of dragonflies and mayflies. Like other corydiid roaches that inhabit xeric areas, Ergaula has an atmospheric water-vapor absorption mechanism that includes a gland and a ductus on the epipharnyx and bladders on the hypopharynx. The mandibular adductor is in cockroaches asymmetric, a pattern not found in termites, mantids, or other closely related insects.

Children who idiopathically toe-walk have greater plantarflexor effective mechanical advantage compared to typically developing children.

PURPOSE: The effective mechanical advantage (EMA) of the plantarflexor muscles is important for gait function and is likely different from typical in equinus gait. However, this has never been quantified for children who idiopathically toe-walk (ITW), despite being routinely altered through clinical intervention. METHODS: This study quantified the Achilles tendon and ground reaction force (GRF) moment arms, and the plantarflexor EMA of 5 children who ITW and 14 typically developing (TD) children, whilst walking on an instrumented treadmill. RESULTS: There was no difference in the Achilles tendon moment arm length throughout stance between groups (p > 0.05). Children who ITW had a significantly greater GRF moment arm length in early stance (20-24% p = 0.001), but a significantly shorter GRF moment arm length during propulsion (68-74% of stance; p = 0.013) than TD children. Therefore, children who ITW had a greater plantarflexor EMA than TD children when active plantarflexion moments were being generated (60-70% of stance; p = 0.007). Consequently, it was estimated that children who ITW required 30% less plantarflexor muscle force for propulsion. CONCLUSION: Clinical decision making should fully consider that interventions which aim to restore a typical heel-toe gait pattern risk compromising this advantageous leverage and thus, may increase the strength requirements for gait.

The diet of early birds based on modern and fossil evidence and a new framework for its reconstruction.

Birds are some of the most diverse organisms on Earth, with species inhabiting a wide variety of niches across every major biome. As such, birds are vital to our understanding of modern ecosystems. Unfortunately, our understanding of the evolutionary history of modern ecosystems is hampered by knowledge gaps in the origin of modern bird diversity and ecosystem ecology. A crucial part of addressing these shortcomings is improving our understanding of the earliest birds, the non-avian avialans (i.e. non-crown birds), particularly of their diet. The diet of non-avian avialans has been a matter of debate, in large part because of the ambiguous qualitative approaches that have been used to reconstruct it. Here we review methods for determining diet in modern and fossil avians (i.e. crown birds) as well as non-avian theropods, and comment on their usefulness when applied to non-avian avialans. We use this to propose a set of comparable, quantitative approaches to ascertain fossil bird diet and on this basis provide a consensus of what we currently know about fossil bird diet. While no single approach can precisely predict diet in birds, each can exclude some diets and narrow the dietary possibilities. We recommend combining (i) dental microwear, (ii) landmark-based muscular reconstruction, (iii) stable isotope geochemistry, (iv) body mass estimations, (v) traditional and/or geometric morphometric analysis, (vi) lever modelling, and (vii) finite element analysis to reconstruct fossil bird diet accurately. Our review provides specific methodologies to implement each approach and discusses complications future researchers should keep in mind. We note that current forms of assessment of dental mesowear, skull traditional morphometrics, geometric morphometrics, and certain stable isotope systems have yet to be proven effective at discerning fossil bird diet. On this basis we report the current state of knowledge of non-avian avialan diet which remains very incomplete. The ancestral dietary condition in non-avian avialans remains unclear due to scarce data and contradictory evidence in Archaeopteryx. Among early non-avian pygostylians, Confuciusornis has finite element analysis and mechanical advantage evidence pointing to herbivory, whilst Sapeornis only has mechanical advantage evidence indicating granivory, agreeing with fossilised ingested material known for this taxon. The enantiornithine ornithothoracine Shenqiornis has mechanical advantage and pedal morphometric evidence pointing to carnivory. In the hongshanornithid ornithuromorph Hongshanornis only mechanical advantage evidence indicates granivory, but this agrees with evidence of gastrolith ingestion in this taxon. Mechanical advantage and ingested fish support carnivory in the songlingornithid ornithuromorph Yanornis. Due to the sparsity of robust dietary assignments, no clear trends in non-avian avialan dietary evolution have yet emerged. Dietary diversity seems to increase through time, but this is a preservational bias associated with a predominance of data from the Early Cretaceous Jehol Lagerstätte. With this new framework and our synthesis of the current knowledge of non-avian avialan diet, we expect dietary knowledge and evolutionary trends to become much clearer in the coming years, especially as fossils from other locations and climates are found. This will allow for a deeper and more robust understanding of the role birds played in Mesozoic ecosystems and how this developed into their pivotal role in modern ecosystems.

The evolutionary biomechanics of locomotor function in giant land animals.

Giant land vertebrates have evolved more than 30 times, notably in dinosaurs and mammals. The evolutionary and biomechanical perspectives considered here unify data from extant and extinct species, assessing current theory regarding how the locomotor biomechanics of giants has evolved. In terrestrial tetrapods, isometric and allometric scaling patterns of bones are evident throughout evolutionary history, reflecting general trends and lineage-specific divergences as animals evolve giant size. Added to data on the scaling of other supportive tissues and neuromuscular control, these patterns illuminate how lineages of giant tetrapods each evolved into robust forms adapted to the constraints of gigantism, but with some morphological variation. Insights from scaling of the leverage of limbs and trends in maximal speed reinforce the idea that, beyond 100-300 kg of body mass, tetrapods reduce their locomotor abilities, and eventually may lose entire behaviours such as galloping or even running. Compared with prehistory, extant megafaunas are depauperate in diversity and morphological disparity; therefore, turning to the fossil record can tell us more about the evolutionary biomechanics of giant tetrapods. Interspecific variation and uncertainty about unknown aspects of form and function in living and extinct taxa still render it impossible to use first principles of theoretical biomechanics to tightly bound the limits of gigantism. Yet sauropod dinosaurs demonstrate that >50 tonne masses repeatedly evolved, with body plans quite different from those of mammalian giants. Considering the largest bipedal dinosaurs, and the disparity in locomotor function of modern megafauna, this shows that even in terrestrial giants there is flexibility allowing divergent locomotor specialisations.

Cheliceral chelal design in free-living astigmatid mites.

Cheliceral chelal design in free-living astigmatid mites (Arthropoda: Acari) is reviewed within a mechanical model. Trophic access (body size and cheliceral reach) and food morsel handling (chelal gape and estimated static adductive crushing force) are morphologically investigated. Forty-seven commonly occurring astigmatid mite species from 20 genera (covering the Acaridae, Aeroglyphidae, Carpoglyphidae, Chortoglyphidae, Glycyphagidae, Lardoglyphidae, Pyroglyphidae, Suidasiidae, and Winterschmidtiidae) are categorised into functional groups using heuristics. Conclusions are confirmed with statistical tests and multivariate morphometrics. Despite these saprophagous acarines in general being simple 'shrunken/swollen' versions of each other, clear statistical correlations in the specifics of their mechanical design (cheliceral and chelal scale and general shape) with the type of habitat and food consumed (their 'biome') are found. Using multivariate analyses, macro- and microsaprophagous subtypes are delineated. Relative ratios of sizes on their own are not highly informative of adaptive syndromes. Sympatric resource competition is examined. Evidence for a maximum doubling of approximate body volume within nominal taxa is detected but larger mites are not more 'generalist' feeding types. Two contrasting types of basic 'Bauplan' are found differing in general scale: (i) a large, chunk-crunching, 'demolition'-feeding omnivore design (comprising 10 macrosaprophagous astigmatid species), and (ii) a small selective picking, squashing/slicing or fragmentary/'plankton' feeding design (which may indicate obligate fungivory/microbivory) comprising 20 microsaprophagous acarid-shaped species. Seventeen other species appear to be specialists. Eleven of these are either: small (interstitial/burrowing) omnivores-or a derived form designed for processing large hard food morsels (debris durophagy, typified by the pyroglyphid Dermatophagoides farinae), or a specialist sub-type of particular surface gleaning/scraping fragmentary feeding. Six possible other minor specialist gleaning/scraping fragmentary feeders types each comprising one to two species are described. Details of these astigmatid trophic-processing functional groups need field validation and more corroborative comparative enzymology. Chelal velocity ratio in itself is not highly predictive of habitat but with cheliceral aspect ratio (or chelal adductive force) is indicative of life-style. Herbivores and pest species are typified by a predicted large chelal adductive force. Pest species may be 'shredders' derived from protein-seeking necrophages. Carpoglyphus lactis typifies a mite with tweezer-like chelae of very feeble adductive force. It is suggested that possible zoophagy (hypocarnivory) is associated with low chelal adductive force together with a small or large gape depending upon the size of the nematode being consumed. Kuzinia laevis typifies an oophagous durophage. Functional form is correlated with taxonomic position within the Astigmata-pyroglyphids and glycyphagids being distinct from acarids. A synthesis with mesostigmatid and oribatid feeding types is offered together with clarification of terminologies. The chelal lyrifissure in the daintiest chelicerae of these astigmatids is located similar to where the action of the chelal moveable digit folds the cheliceral shaft in uropodoids, suggesting mechanical similarities of function. Acarid astigmatids are trophically structured like microphytophagous/fragmentary feeding oribatids. Some larger astigmatids (Aleuroglyphus ovatus, Kuzinia laevis, Tyroborus lini) approximate, and Neosuidasia sp. matches, the design of macrophytophagous oribatids. Most astigmatid species reviewed appear to be positioned with other oribatid secondary decomposers. Only Dermatophagoides microceras might be a primary decomposer approximating a lichenivorous oribatid (Austrachipteria sp.) in trophic form. Astigmatid differences are consilient with the morphological trend from micro- to macrophytophagy in oribatids. The key competency in these actinotrichid mites is a type of 'gnathosomisation' through increased chelal and cheliceral height (i.e., a shape change that adjusts the chelal input effort arm and input adductive force) unrestricted by the dorsal constraint of a mesostigmatid-like gnathotectum. A predictive nomogram for ecologists to use on field samples is included. Future work is proposed in detail.

Dental microwear texture analysis along reptile tooth rows: complex variation with non-dietary variables.

Dental microwear texture analysis (DMTA) is a powerful technique for reconstructing the diets of extant and extinct taxa. Few studies have investigated intraspecific microwear differences along with tooth rows and the influence of endogenous non-dietary variables on texture characteristics. Sampling teeth that are minimally affected by non-dietary variables is vital for robust dietary reconstructions, especially for taxa with non-occlusal (non-chewing) dentitions as no standardized sampling strategies currently exist. Here, we apply DMTA to 13 species of extant reptile (crocodilians and monitor lizards) to investigate intraspecific microwear differences along with tooth rows and to explore the influence of three non-dietary variables on exhibited differences: (i) tooth position, (ii) mechanical advantage, and (iii) tooth aspect ratio. Five species exhibited intraspecific microwear differences. In several crocodilians, the distally positioned teeth exhibited the 'roughest' textures, and texture characteristics correlated with all non-dietary variables. By contrast, the mesial teeth of the roughneck monitor (Varanus rudicollis) exhibited the 'roughest' textures, and texture characteristics did not correlate with aspect ratio. These results are somewhat consistent with how reptiles preferentially use their teeth during feeding. We argue that DMTA has the potential to track mechanical and behavioural differences in tooth use which should be taken into consideration in future dietary reconstructions.

Rules of nature's Formula Run: Muscle mechanics during late stance is the key to explaining maximum running speed.

The maximum running speed of legged animals is one evident factor for evolutionary selection-for predators and prey. Therefore, it has been studied across the entire size range of animals, from the smallest mites to the largest elephants, and even beyond to extinct dinosaurs. A recent analysis of the relation between animal mass (size) and maximum running speed showed that there seems to be an optimal range of body masses in which the highest terrestrial running speeds occur. However, the conclusion drawn from that analysis-namely, that maximum speed is limited by the fatigue of white muscle fibres in the acceleration of the body mass to some theoretically possible maximum speed-was based on coarse reasoning on metabolic grounds, which neglected important biomechanical factors and basic muscle-metabolic parameters. Here, we propose a generic biomechanical model to investigate the allometry of the maximum speed of legged running. The model incorporates biomechanically important concepts: the ground reaction force being counteracted by air drag, the leg with its gearing of both a muscle into a leg length change and the muscle into the ground reaction force, as well as the maximum muscle contraction velocity, which includes muscle-tendon dynamics, and the muscle inertia-with all of them scaling with body mass. Put together, these concepts' characteristics and their interactions provide a mechanistic explanation for the allometry of maximum legged running speed. This accompanies the offering of an explanation for the empirically found, overall maximum in speed: In animals bigger than a cheetah or pronghorn, the time that any leg-extending muscle needs to settle, starting from being isometric at about midstance, at the concentric contraction speed required for running at highest speeds becomes too long to be attainable within the time period of a leg moving from midstance to lift-off. Based on our biomechanical model, we, thus, suggest considering the overall speed maximum to indicate muscle inertia being functionally significant in animal locomotion. Furthermore, the model renders possible insights into biological design principles such as differences in the leg concept between cats and spiders, and the relevance of multi-leg (mammals: four, insects: six, spiders: eight) body designs and emerging gaits. Moreover, we expose a completely new consideration regarding the muscles' metabolic energy consumption, both during acceleration to maximum speed and in steady-state locomotion.

A review of linkage mechanisms in animal joints and related bioinspired designs.

This paper presents a review of biological mechanical linkage mechanisms. One purpose is to identify the range of kinematic functions that they are able to perform. A second purpose is to review progress in bioinspired designs. Ten different linkage mechanisms are presented. They are chosen because they cover a wide range of functionality and because they have potential for bioinspired design. Linkage mechanisms enable animal joints to perform highly sophisticated and optimised motions. A key function of animal linkage mechanisms is the optimisation of actuator location and mechanical advantage. This is crucially important for animals where space is highly constrained. Many of the design features used by engineers in linkage mechanisms are seen in nature, such as short coupler links, extended bars, elastic energy storage and latch mechanisms. However, animal joints contain some features rarely seen in engineering such as integrated cam and linkage mechanisms, nonplanar four-bar mechanisms, resonant hinges and highly redundant actuators. The extreme performance of animal joints together with the unusual design features makes them an important area of investigation for bioinspired designs. Whilst there has been significant progress in bioinspiration, there is the potential for more, especially in robotics where compactness is a key design driver.

The scaling of ground reaction forces and duty factor in monitor lizards: implications for locomotion in sprawling tetrapods.

Geometric scaling predicts a major challenge for legged, terrestrial locomotion. Locomotor support requirements scale identically with body mass (α M1), while force-generation capacity should scale α M2/3 as it depends on muscle cross-sectional area. Mammals compensate with more upright limb postures at larger sizes, but it remains unknown how sprawling tetrapods deal with this challenge. Varanid lizards are an ideal group to address this question because they cover an enormous body size range while maintaining a similar bent-limb posture and body proportions. This study reports the scaling of ground reaction forces and duty factor for varanid lizards ranging from 7 g to 37 kg. Impulses (force×time) (α M0.99-1.34) and peak forces (α M0.73-1.00) scaled higher than expected. Duty factor scaled α M0.04 and was higher for the hindlimb than the forelimb. The proportion of vertical impulse to total impulse increased with body size, and impulses decreased while peak forces increased with speed.

Mechanical advantage and joint function of the lower limb during hopping at different frequencies.

Mechanical output at a joint level could be influenced by its leverage characteristics and by its functional behaviour and both could change to accommodate the demands of a given locomotor task. In this study, the mechanical power generated at the knee and ankle joints and their functional indexes (i.e. damper, strut, spring and motor like-function) were calculated by using 3D kinematic and kinetic data during hopping at 2, 2.5, 3 and 3.5 Hz. The effective mechanical advantage (i.e. the ratio between internal and external moment arm) of the knee (EMAK) and ankle (EMAA) and joint stiffness were calculated as well. Joint stiffness increased with frequency whereas positive and negative joint power decreased with it, the ankle power values being always larger (20-50%) than those at the knee. EMAA reached its highest value (0.4) during the propulsive phase at 3 Hz whereas no significant changes in EMAK were observed as a function of frequency in both the absorption and propulsive phases. Knee joint-functional index shifted from a spring to a strut-like function with increasing frequency (from 56 to 8% and from 4 to 51%, respectively) while the ankle operated mainly as a spring (from 90 to 53%), its damper and motor-like indexes being negligible at all frequencies (<5%). Therefore, in hopping, the knee works to dissipate mechanical energy (the combination of its damper and strut indexes increase from 23 to 72% at these frequencies) and the primary source of mechanical power is attributable to the elastic function of the ankle.

Insectivory leads to functional convergence in a group of Neotropical rodents.

The mandible of vertebrates serves as insertion area for masticatory muscles that originate on the skull, and its functional properties are subject to selective forces related to trophic ecology. The efficiency of masticatory muscles can be measured as mechanical advantage on the mandible, which, in turn, has the property of correlating with bite force and shape. In the present work, we quantify the mechanical advantage of the mandible of akodontine rodents, which present a diverse radiation of insectivorous specialists, to assess their relationship to the estimated bite force and diet. We also tested the degree of morphofunctional convergence in response to insectivory on the group. We found the mechanical advantages to be convergent on insectivorous species, and associated with the estimated bite force, with higher mechanical advantages in species with a stronger bite and short, robust mandibles and lower mechanical advantages in insectivorous species with weaker bites and more elongated, dorso-ventrally compressed mandibles. Insectivorous species of Akodontini are functional specialists for the consumption of live prey and may exploit the resources that shrews, moles and hedgehogs consume elsewhere.

Do Tangential Finger Forces Utilize Mechanical Advantage During Moment of Force production?

This study investigated the beneficial effects of the utilization of mechanical advantage (MA) of finger tangential forces during the moment production. Subjects produced the resistive moment of force against the external torque while the moment arms of the tangential forces were systematically changed. We observed a relatively large contribution to the net moment by the tangential forces with the increased moment arms, whereas the vector sum of normal and tangential forces decreased. The indices of multi-finger coordination for the stabilization of the moment of forces and force direction increased with the moment arms. The current results provide evidence that the utilization of MA is associated with both the efficiency of force production and the stabilization of performance variables.

Comparable behaviour of ring and little fingers due to an artificial reduction in thumb contribution to hold objects.

BACKGROUND: The human hand plays a crucial role in accomplishing activities of daily living. The contribution of each finger in the human hand is remarkably unique in establishing object stabilization. According to the mechanical advantage hypothesis, the little finger tends to exert a greater normal force than the ring finger during a supination moment production task to stabilize the object. Similarly, during pronation, the index finger produces more normal force when compared with the middle finger. Hence, the central nervous system employs the peripheral fingers for torque generation to establish the equilibrium as they have a mechanical advantage of longer moment arms for normal force. In our study, we tested whether the mechanical advantage hypothesis is supported in a task in which the contribution of thumb was artificially reduced. We also computed the safety margin of the individual fingers and thumb. METHODOLOGY: Fifteen participants used five-finger prismatic precision grip to hold a custom-built handle with a vertical railing on the thumb side. A slider platform was placed on the railing such that the thumb sensor could move either up or down. There were two experimental conditions. In the "Fixed" condition, the slider was mechanically fixed, and hence the thumb sensor could not move. In the "Free" condition, the slider platform on which the thumb sensor was placed could freely move. In both conditions, the instruction was to grasp and hold the handle (and the platform) in static equilibrium. We recorded tangential and normal forces of all the fingers. RESULTS: The distribution of fingertip forces and moments changed depending on whether the thumb platform was movable (or not). In the free condition, the drop in the tangential force of thumb was counteracted by an increase in the normal force of the ring and little finger. Critically, the normal forces of the ring and little finger were statistically equivalent. The safety margin of the index and middle finger did not show a significant drop in the free condition when compared to fixed condition. CONCLUSION: We conclude that our results does not support the mechanical advantage hypothesis at least for the specific mechanical task considered in our study. In the free condition, the normal force of little finger was comparable to the normal force of the ring finger. Also, the safety margin of the thumb and ring finger increased to prevent slipping of the thumb platform and to maintain the handle in static equilibrium during the free condition. However, the rise in the safety margin of the ring finger was not compensated by a drop in the safety margin of the index and middle finger.

Differences in muscle mechanics underlie divergent optimality criteria between feeding and locomotor systems.

Tetrapod musculoskeletal diversity is usually studied separately in feeding and locomotor systems. However, direct 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. Recent studies using this approach have proposed that the feeding system is optimized for precise application of high forces and the locomotor system is optimized for wide and rapid joint excursions for minimal energetic expenditure. If this hypothesis is correct, then it stands to reason that other anatomical and biomechanical variables within the feeding and locomotor systems should reflect these diverging functions. To test this hypothesis, we compared muscle moment arm lengths, mechanical advantages, and force vector orientations of two jaw elevator muscles (m. temporalis and m. superficial masseter), an elbow flexor (m. brachialis) and extensor (m. triceps- lateral head), and a knee flexor (m. biceps femoris-short head) and extensor (m. vastus lateralis) across 18 species of primates. Our results show that muscles of the feeding system are more orthogonally oriented relative to the resistance arm (mandible) and operate at relatively large moment arms and mechanical advantages. Moreover, these variables show relatively little change across the range of jaw excursion. In contrast, the representative muscles of the locomotor system have much smaller mechanical advantages and, depending on joint position, smaller muscle moment arm lengths and almost parallel orientations relative to the resistance arm. These patterns are consistent regardless of phylogeny, body mass, locomotor mode, and feeding specialization. We argue that these findings reflect fundamental functional dichotomies between tetrapod locomotor and feeding systems. By organizing muscles in a manner such that moment arms and mechanical advantage are relatively small, the locomotor system can produce broad joint excursions and high angular velocities with only small muscular contraction. As such, the anatomical organization of muscles within the limbs allows striding animals to move relatively rapidly and with minimal energetic expenditure. In contrast, the anatomical configuration of muscles in the feeding system, at least m. superficial masseter and m. temporalis, favors their force-producing capacity at the expense of excursion and velocity.

Effective Mechanical Advantage About the Ankle Joint and the Effect of Achilles Tendon Curvature During Toe-Walking.

Aim: To study the causes of locomotor dysfunction, estimate muscle forces, or understand the influence of altered sarcomere and muscle properties and behaviours on whole body function, it is necessary to examine the leverage with which contractile forces operate. At the ankle joint, current methods to quantify this leverage for the plantarflexors do not account for curvature of the Achilles tendon, and so may not be appropriate when studying equinus gait. Thus, novel methodologies need to be developed and implemented to quantify the Achilles tendon moment arm length during locomotion. Methods: Plantarflexor internal moment arm length and effective mechanical advantage of 11 typically developed young adults were calculated throughout stance, while heel-toe walking and voluntarily toe-walking on an instrumented treadmill. Achilles tendon moment arm was defined in two-ways: (1) assuming a straight tendon, defined between the gastrocnemius medialis myotendinous junction and Achilles tendon insertion point, and (2) accounting for tendon curvature, by tracking the initial path of the Achilles tendon from the calcaneal insertion. Results: When accounting for tendon curvature, Achilles tendon moment arm length and plantarflexor effective mechanical advantage did not differ between walking conditions (p > 0.05). In contrast, when assuming a straight tendon, Achilles tendon moment arm length (p = 0.043) and plantarflexor effective mechanical advantage (p = 0.007) were significantly greater when voluntary toe-walking than heel-toe walking in late stance. Discussion: Assuming a straight Achilles tendon led to a greater Achilles tendon moment arm length and plantarflexor effective mechanical advantage during late stance, compared to accounting for tendon curvature. Consequently, plantarflexor muscle force would appear smaller when assuming a straight tendon. This could lead to erroneous interpretations of muscular function and fascicle force-length-velocity behaviour in vivo, and potentially inappropriate and ineffective clinical interventions for equinus gait.

Leverage mechanical alterations during walking at self-selected speed in patients with Parkinson's disease.

Individuals with Parkinson's disease (PD) show poor walking performance compared to healthy adults. Leverage changes may provide insight into this walking abnormality, since they have important effects on both biomechanical and physiological variables. Hence, we investigated the differences in internal and external moment arms at the knee and ankle joints, as well as the effective mechanical advantage during walking at self-selected speed. Furthermore, the effects on walking of a simultaneous cognitive task were analysed. Kinetic (resultant ground reaction force and joint moments), kinematic (movement speed) and mechanical leverage (internal and external moment arms) parameters of 10 mild-to-moderate PD patients and 10 age-matched controls were measured in single and dual task condition. Finally, effective mechanical advantage was calculated as the ratio between internal and external moment arm for each joint. PD patients had a slower walking and showed larger and lower values of knee and ankle joint moments, respectively. No difference in force among groups was recorded. External moment arms were larger (in both joints) for PD, whereas slight changes were observed for internal moment arms. Consequently, effective mechanical advantage values seemed to be lower for PD. Surprisingly, leverage difference among groups was reduced during the dual task condition, resulting in a "more effective" walking strategy for PD. These findings suggest that during single task PD patients have several leverage disadvantages, which could affect the joint assessment. On the contrary, during dual task they reduced these mechanical negative effects by positively obtaining normal values of effective mechanical advantage.

Scaling of Anatomically Derived Maximal Bite Force in Primates.

By combining muscle architectural data with biomechanical variables relating to the jaw, we produce anatomically derived maximum bite force estimations for 23 species of catarrhine and platyrrhine primates. We investigate how bite force scales across the sample as a whole (and within each parvorder) relative to two size proxies, body mass and cranial geometric mean, and the effect of diet upon bite force. Bite force is estimated at three representative bite points along the dental row: the first maxillary incisor, canine, and third-most mesial paracone. We modeled bite force by combining calculated physiological cross-sectional area of the jaw adductors from Hartstone-Rose et al. [Anat Rec 301 (2018) 311-324] with osteological measurements of lever- and load-arm lengths from the same specimens [Hartstone-Rose et al., Anat Rec 295 (2012) 1336-1351]. Bite force scales with positive allometry relative to cranial geometric mean across our entire sample and tends toward positive allometry relative to body mass. Bite force tends toward positive allometry within platyrrhines but scales isometrically within catarrhines. There was no statistically significant scaling difference with diet. Our findings imply an absence of a dietary signal in the scaling of bite force, a result that differs from the scaling of physiological cross-sectional area alone. That is, although previous studies have found a dietary signal in the muscle fiber architecture in these species, when these are combined with their leverages, that signal is undetectable. On the parvorder level, our data also demonstrate that the platyrrhine masticatory system appears more mechanically advantageous than that of catarrhines. Anat Rec, 2019. © 2019 American Association for Anatomy Anat Rec, 303:2026-2035, 2020. © 2019 American Association for Anatomy.