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The mealworm Tenebrio molitor L. (Coleoptera: Tenebrionidae) feeds on wheat bran and is considered both a pest and an edible insect. Its larvae contain proteins and essential amino acids, fats, and minerals, making them suitable for animal and human consumption. Zearalenone (ZEA) is the mycotoxin most commonly associated with Fusarium spp. It is found in cereals and cereal products, so their consumption is a major risk for mycotoxin contamination. One of the most important effects of ZEA is the induction of oxidative stress, which leads to physiological and behavioral changes. This study deals with the effects of high doses of ZEA (10 and 20 mg/kg) on survival, molting, growth, weight gain, activity of antioxidant enzymes superoxide dismutase (SOD) and glutathione S-transferase (GST), and locomotion of mealworm larvae. Both doses of ZEA were found to (i) have no effect on survival, (ii) increase molting frequency, SOD, and GST activity, and (iii) decrease body weight and locomotion, with more pronounced changes at 20 mg/kg. These results indicated the susceptibility of T. molitor larvae to high doses of ZEA in feed. [ABSTRACT FROM AUTHOR]

Small vertebrates, such as geckos, often use their tails to regulate their posture after take-off and suppress abnormal rotations in mid-air (such as sliding and falling). However, research on tail function before take-off remains scarce, particularly when animals overcome gravity to jump upward. This study examined tailed and tailless geckos' ability to jump over obstacles. The tails swung up and reached their upper limits during take-off, corresponding to a continuous increase in their trunk elevation angle. Compared to tailless geckos, geckos with intact tails had lower body and tail angular velocities. In addition, tailed geckos achieved larger take-off angles and higher jumps; moreover, their take-off angles were consistent with their velocity angles. Theoretical analyses demonstrated that the tail and its upward swing before take-off could benefit lifting the forelimb and hindlimbs off the ground, expanding the stability area of their centre of mass (COM). A robot prototype verified the effect of the tail and its swing on trunk elevation before take-off. Other parameters, such as the posture of the hindlimbs (support length and angle) and initial angle of the trunk, affect posture regulation. This study significantly extends our knowledge in animal locomotion and provides indispensable information for improving robot jumping ability. [ABSTRACT FROM AUTHOR]

While several marine species exhibit biological rhythms of heart rate, gill ventilation, or locomotion, the relationship between these three measures in any species remains unexplored. The American horseshoe crab, Limulus polyphemus Linnaeus, 1758 , expresses circalunidian locomotor rhythms and circadian eye sensitivity rhythms, but it is not clear if either heart and ventilation rates are controlled on a circadian, or circatidal basis or the nature of the relationship between these three measures. We aimed to determine the extent to which the heart and ventilation rates of L. polyphemus are coordinated with its endogenous rhythms of locomotion. We recorded rhythmic beating of the heart and movements of the gill plates in freely moving horseshoe crabs. Most individuals exhibited robust circatidal rhythms of locomotion, heart rate, and ventilation that were highly correlated with each other over three weeks of continuous recording. These results are the first showing long-term rhythms of all three measures in any marine species and suggest that heart rate and ventilation rhythms are coordinated in L. polyphemus , both with each other, and with locomotion, and thus are all modulated on a tidal basis. [ABSTRACT FROM AUTHOR]

Scaling of body shape, gait, limb kinematics, and kinetics can inform about mechanical and energetic constraints influencing animal locomotion on land. However, our knowledge of terrestrial locomotion among tetrapods is biased toward animals that use parasagittal limb postures (mammals and birds). Whether species that use non-parasagittal limb postures (e.g. crocodylians and lizards) share similar scaling principles remains uncertain, especially toward the larger end of the body size spectrum. Here, we compare new data from large adult American alligators (140–223 kg) with previous data from smaller individuals (0.2–5.6 kg) to evaluate the scaling of gait, limb posture, ground reaction forces, and inertial properties of body segments over a size range spanning three orders of magnitude. Larger alligators walked with higher relative stride frequencies and shorter relative stride lengths at equal dimensionless speeds, suggesting that alligators prioritize stance phase mechanical work reduction at the expense of higher costs for swing as they increase in size. Limb posture also changed with body mass: ∼1 kg and larger alligators typically used more adducted and extended limbs, which is expected to mitigate limb muscle forces during stance. Ground reaction forces were increasingly hindlimb dominant as alligators grow to larger size, paralleled by a posterior shift in the center of mass. Counterintuitively, the forelimb grew faster in mass than the hindlimb, possibly because recruited muscle volume is closer to its maximum capacity in the forelimb, compelling larger alligators to keep up with increased mechanical demands via muscle growth more for the forelimb than the hindlimb. These results highlight the numerous changes in body proportions and locomotor performance that are required among animals that use non-parasagittal limb posture, as they grow from neonatal sizes into large adults. [ABSTRACT FROM AUTHOR]

Birds are well known for their ability to fly, and flight-capable adult birds have many anatomical specializations for meeting the demands of aerial locomotion. Juvenile birds in altricial species typically acquire these specializations close to fledging and leave the nest with some flight capability. In contrast, juveniles in most precocial species begin navigating their environment with rudimentary anatomies and may not develop full-sized wings or musculoskeletal apparatuses for several months. This manuscript explores how juvenile birds achieve high levels of locomotor performance in the absence of flight specializations, by synthesizing work on two groups of precocial birds with very different developmental strategies. Galliforms like the Chukar Partridge (Alectoris chukar) have early wing development and are capable of flight within weeks. Compared with adults, juvenile chukars have less aerodynamically effective feathers and smaller muscles but compensate through anatomical, kinematic, and behavioral mechanisms. In contrast, waterfowl have delayed wing development and initially rely on leg-based locomotion. In Mallards (Anas platyrhynchos) and their domesticated derivatives, leg investment and performance peak early in ontogeny, but then decline when wings develop. Chukar and mallard juveniles thus rely on different mechanisms for negotiating their surroundings in the absence of flight specializations. In conjunction with work in other animals, these patterns indicate that juveniles with developing locomotor apparatuses can achieve surprisingly high levels of locomotor performance through a variety of compensatory mechanisms. [ABSTRACT FROM AUTHOR]

While animals swim, crawl, walk, and fly with apparent ease, building robots capable of robust locomotion remains a significant challenge. In this review, we draw attention to mechanosensation—the sensing of mechanical forces generated within and outside the body—as a key sense that enables robust locomotion in animals. We discuss differences between mechanosensation in animals and current robots with respect to (1) the encoding properties and distribution of mechanosensors and (2) the integration and regulation of mechanosensory feedback. We argue that robotics would benefit greatly from a detailed understanding of these aspects in animals. To that end, we highlight promising experimental and engineering approaches to study mechanosensation, emphasizing the mutual benefits for biologists and engineers that emerge from moving forward together. [ABSTRACT FROM AUTHOR]

Locomotion triggers a coordinated response of both neurons and astrocytes in the brain. Here we performed calcium (Ca2+) imaging of these two cell types in the somatosensory cortex in head-fixed mice moving on the airlifted platform. Ca2+ activity in astrocytes significantly increased during locomotion from a low quiescence level. Ca2+ signals first appeared in the distal processes and then propagated to astrocytic somata, where it became significantly larger and exhibited oscillatory behaviour. Thus, astrocytic soma operates as both integrator and amplifier of Ca2+ signal. In neurons, Ca2+ activity was pronounced in quiescent periods and further increased during locomotion. Neuronal Ca2+ concentration ([Ca2+]i) rose almost immediately following the onset of locomotion, whereas astrocytic Ca2+ signals lagged by several seconds. Such a long lag suggests that astrocytic [Ca2+]i elevations are unlikely to be triggered by the activity of synapses among local neurons. Ca2+ responses to pairs of consecutive episodes of locomotion did not significantly differ in neurons, while were significantly diminished in response to the second locomotion in astrocytes. Such astrocytic refractoriness may arise from distinct mechanisms underlying Ca2+ signal generation. In neurons, the bulk of Ca2+ enters through the Ca2+ channels in the plasma membrane allowing for steady-level Ca2+ elevations in repetitive runs. Astrocytic Ca2+ responses originate from the intracellular stores, the depletion of which affects subsequent Ca2+ signals. Functionally, neuronal Ca2+ response reflects sensory input processed by neurons. Astrocytic Ca2+ dynamics is likely to provide metabolic and homeostatic support within the brain active milieu. Graphical Abstract [ABSTRACT FROM AUTHOR]

The mangrove rivulus (Kryptolebias marmoratus) is a phenotypically plastic teleost fish that can spend considerable time on land and traverse the terrestrial realm through a behavior termed the tail-flip jump. The tail-flip jump is a transitional stage between fully aquatic and terrestrial lifestyles. Therefore, understanding this behavior can provide insight into how organisms adapt to new environments over evolutionary time. Studies of K. marmoratus show that terrestrial acclimation and exercise improve tail-flip jumping performance due to muscle remodeling, but the implications of these muscular changes on aquatic locomotion are unknown. In the present study, we hypothesized that (1) terrestrial acclimation and exercise lead to physiological changes, such as changes to muscle fiber type, muscle mass distribution, or body shape, that optimize tail-flip jump distance and endurance while negatively impacting swimming performance in K. marmoratus , and (2) plasticity of the brain (which has been demonstrated in response to a variety of stimuli in K. marmoratus) allows terrestrial emersion and exercise to cause behavioral changes that promote survival and long-term reproductive success. To test these hypotheses, we measured the critical swimming speed (U crit), tail-flip jump distance, terrestrial endurance, and undisturbed aquatic behavior of age- and size-matched K. marmoratus before and after a terrestrial exercise period. This period consisted of six 3-min exercise sessions spread over 12 days, during which the fish were prompted to jump continuously. To isolate the effects of air exposure, a separate group was exposed to air for an equivalent period but not allowed to jump. Air exposure improved maximum jump distance but negatively affected swimming performance (U crit). Terrestrial endurance (number of jumps) improved in the exercised group, but U crit showed no significant change. Contrary to our first hypothesis, a trade-off exists between jump distance and U crit but not between jump endurance and U crit. Exercised individuals were more active following exercise, resulting either from the onset of dispersion behavior or a heightened stress response. [ABSTRACT FROM AUTHOR]

In tetrapods, BMP-signaling coordinates limb outgrowth, skeleton patterning, and apoptosis during the formation of their typical autopod structures, the digits. In addition, inhibition of BMP signaling during mouse limb development leads to the persistence and enlargement of an important signaling center, the apical ectodermal ridge (AER), and consequent digit defects. Interestingly, during fish fin development there is a natural elongation of the AER, rapidly converted into an apical finfold (FF), in which osteoblasts differentiate into dermal fin-rays used in aquatic locomotion. Previous reports have led us to suggest that upregulation of Hox13 genes in the distal fin mesenchyme, caused by the origin of novel enhancer modules, may have caused an increment of the BMP signaling potentially leading to the apoptosis of these osteoblasts precursors of the fin-rays. To explore this hypothesis, we characterized the expression of several components of the BMP signaling in zebrafish lines with distinct FF sizes (bmp2b , smad1 , smoc1 , smoc2, grem1a, msx1b, msx2b, Psamd1/5/9). Our data suggest that the BMP signaling is enhanced in shorter FFs and inhibited in longer FFs, as implied by the differential expression of several components of this network. In addition, we detected an earlier expression of several of these BMP-signaling components associated with the development of short FFs and the opposite tendency during the development of longer FFs. Thus, our results suggest that a heterochronic shift, involving enhanced Hox13 expression and BMP signaling may have caused the reduction of the FF size during the evolutionary transition from fish fins to tetrapod limbs. [ABSTRACT FROM AUTHOR]

Biological variation is often considered in a scalable hierarchy, e.g. within the individual, within the populations, above the species level. Morphological integration, the concept of covariation among constituent parts of an organism, is also hierarchical; the degree to which these "modules" covary is a matter of the scale of the study as well as underlying processes driving the covariation. Multilevel analyses of trait covariation are a valuable tool to infer the origins and historical persistence of morphological diversity. Here, we investigate concordance in patterns of integration and modularity across three biological levels of variation: within a species, within two genera-level radiations, and among species at the family level. We demonstrate this approach using the skull of mammalian family Leporidae (rabbits and hares), which is morphologically diverse and has a rare-among-mammals functional signal of locomotion adaptation. We tested three alternative hypotheses of modularity; from the most supported we investigated disparity and integration of each module to infer which is most responsible for patterns of cranial variation across these levels, and whether variation is partitioned consistently across levels. We found a common pattern of modularity underlies leporid cranial diversity, though there is inconsistency across levels in each module's disparity and integration. The face module contributes the most to disparity at all levels, which we propose is facilitating evolutionary diversity in this clade. Therefore, the distinctive facial tilt of leporids is an adaptation to locomotory behavior facilitated by a modular system that allows lineages to respond differently to selection pressures. [ABSTRACT FROM AUTHOR]

Because of pleiotropy, mutations affect the expression and inheritance of multiple traits and, together with selection, are expected to shape standing genetic covariances between traits and eventual phenotypic divergence between populations. It is therefore important to find if the M matrix, describing mutational variances of each trait and covariances between traits, varies between genotypes. We here estimate the M matrix for six locomotion behavior traits in lines of two genotypes of the nematode Caenorhabditis elegans that accumulated mutations in a nearly neutral manner for 250 generations. We find significant mutational variance along at least one phenotypic dimension of the M matrices, but neither their size nor their orientation had detectable differences between genotypes. The number of generations of mutation accumulation, or the number of MA lines measured, was likely insufficient to sample enough mutations and detect potentially small differences between the two M matrices. We then tested if the M matrices were similar to one G matrix describing the standing genetic (co)variances of a population derived by the hybridization of several genotypes, including the two measured for M, and domesticated to a lab-defined environment for 140 generations. We found that the M and G were different because the genetic covariances caused by mutational pleiotropy in the two genotypes are smaller than those caused by linkage disequilibrium in the lab population. We further show that M matrices differed in their alignment with the lab population G matrix. If generalized to other founder genotypes of the lab population, these observations indicate that selection does not shape the evolution of the M matrix for locomotion behavior in the short-term of a few tens to hundreds of generations and suggests that the hybridization of C. elegans genotypes allows selection on new phenotypic dimensions of locomotion behavior. [ABSTRACT FROM AUTHOR]

Understanding the effect of body size on flight costs is critical for the development of models of aerodynamics and animal energetics. Prior scaling studies that have shown that flight costs scale hypometrically have focused primarily on larger (>100 mg) insects and birds, but most flying species are smaller. We studied the flight physiology of 13 stingless bee species over a large range of body sizes (1–115 mg). Metabolic rate during hovering scaled hypermetrically (scaling slope = 2.11). Larger bees had warm thoraxes, while small bees were nearly ecothermic; however, even controlling for body temperature variation, flight metabolic rate scaled hypermetrically across this clade. Despite having a lower mass-specific metabolic rate during flight, smaller bees could carry the same proportional load. Wingbeat frequency did not vary with body size, in contrast to most studies that find wingbeat frequency increases as body size decreases. Smaller stingless bees have a greater relative forewing surface area, which may help them reduce the energy requirements needed to fly. Further, we hypothesize that the relatively larger heads of smaller species may change their body pitch in flight. Synthesizing across all flying insects, we demonstrate that the scaling of flight metabolic rate changes from hypermetric to hypometric at ∼58 mg body mass with hypermetic scaling below (slope = 1.2) and hypometric scaling (slope = 0.67) >58 mg in body mass. The reduced cost of flight likely provides selective advantages for the evolution of small body size in insects. The biphasic scaling of flight metabolic rates and wingbeat frequencies in insects supports the hypothesis that the scaling of metabolic rate is closely related to the power requirements of locomotion and cycle frequencies. [ABSTRACT FROM AUTHOR]

A "walking" gait has been identified in a range of vertebrate species with different body plans, habitats, and life histories. With increased application of this broad umbrella term, it has become necessary to assess the physical characteristics, analytical approaches, definitions, and diction used to describe walks. To do this, we reviewed studies of slow-speed locomotion across a range of vertebrates to refine the parameters used to define walking, evaluate analytical techniques, and propose approaches to maximize consistency across subdisciplines of zoology. We summarize nine key parameters used to characterize walking behaviors in mammals, birds, reptiles, amphibians, and fish. After identifying consistent patterns across groups, we propose a comprehensive definition for a walking gait. A walk is a form of locomotion where the majority of the forward propulsion of the animal comes from forces generated by the appendages interacting with the ground. During a walk, an appendage must be out of phase with the opposing limb in the same girdle and there is always at least one limb acting as ground-support (no suspension phase). Additionally, walking occurs at dimensionless speeds <1 v * and the average duty factor of the limbs is always >0.5. Relative to other gaits used by the same species, the stance duration of a walk is long, the cycle frequency is low, and the cycle distance is small. Unfortunately, some of these biomechanical parameters, while effectively describing walks, may also characterize other, non-walking gaits. Inconsistent methodology likely contributes to difficulties in comparing data across many groups of animals; consistent application of data collection and analytical techniques in research methodology can improve these comparisons. Finally, we note that the kinetics of quadrupedal movements are still poorly understood and much work remains to be done to understand the movements of small, ecothermic tetrapods. [ABSTRACT FROM AUTHOR]

Many ecological factors influence animal movement, including properties of the media that they move on or through. Animals moving in terrestrial environments encounter conditions that can be challenging for generating propulsion and maintaining stability, such as inclines and deformable substrates that can cause slipping and sinking. In response, tetrapods tend to adopt a more crouched posture and lower their center of mass on inclines and increase the surface area of contact on deformable substrates, such as sand. Many amphibious fishes encounter the same challenges when moving on land, but how these finned animals modulate their locomotion with respect to different environmental conditions and how these modifications compare with those seen within tetrapods is relatively understudied. Mudskippers (Gobiidae: Oxudercinae) are a particularly noteworthy group of amphibious fishes in this context given that they navigate a wide range of environmental conditions, from flat mud to inclined mangrove trees. They use a unique form of terrestrial locomotion called "crutching," where their pectoral fins synchronously lift and vault the front half of the body forward before landing on their pelvic fins, while the lower half of the body and tail are kept straight. However, recent work has shown that mudskippers modify some aspects of their locomotion when crutching on deformable surfaces, particularly those at an incline. For example, on inclined dry sand, mudskippers bent their bodies laterally and curled and extended their tails to potentially act as a secondary propulsor and/or anti-slip device. In order to gain a more comprehensive understanding of the functional diversity and context-dependency of mudskipper crutching, we compared their kinematics on different combinations of substrate types (solid, mud, and dry sand) and inclines (0°, 10°, and 20°). In addition to increasing lateral bending on deformable and inclined substrates, we found that mudskippers increased the relative contact time and contact area of their paired fins, while becoming more crouched, which are responses comparable to those seen in tetrapods and other amphibious fish. Mudskippers on these substrates also exhibited previously undocumented behaviors, such as extending and adpressing the distal portions of their pectoral fins more anteriorly, dorsoventrally bending their trunk, "belly-flopping" on sand, and "gripping" the mud substrate with their pectoral fin rays. Our study highlights potential compensatory mechanisms shared among vertebrates in terrestrial environments while also illustrating that locomotor flexibility and even novelty can emerge when animals are challenged with environmental variation. [ABSTRACT FROM AUTHOR]

The spring-mass model is a model of locomotion aimed at giving the essential mathematical laws of the trajectory of the center of mass of an animal during bouncing gaits, such as hopping (one-dimensional) and running (two-dimensional). This reductionist mechanical system has been extensively investigated for locomotion over horizontal surfaces, whereas it has been largely neglected on other ecologically relevant surfaces, including inclines. For example, how the degree of inclination impacts the dynamics of the center of mass of the spring-mass model has not been investigated thoroughly. In this work, we derive a mathematical model which extends the spring-mass model to inclined surfaces. Among our results, we derive an approximate solution of the system, assuming a small angular sweep of the limb and a small spring compression during stance, and show that this approximation is very accurate, especially for small inclinations of the ground. Furthermore, we derive theoretical bounds on the difference between the Lagrangian and Lagrange equations of the true and approximate systems, and discuss locomotor stability questions of the approximate solutions. We test our models through a sensitivity analysis using parameters relevant to the locomotion of bipedal animals (quail, pheasant, guinea fowl, turkey, ostrich, and humans) and compare our approximate solution to the numerically derived solution of the exact system. We compare the two-dimensional spring-mass model on inclines with the one-dimensional spring-mass model to which it reduces under the limit of no horizontal velocity; we compare the two-dimensional spring-mass model on inclines with the inverted pendulum model on inclines towards which it converges in the case of high stiffness-to-mass ratio. We include comparisons with historically prevalent no-gravity approximations of these models, as well. The insights we have gleaned through all these comparisons and the ability of our approximation to replicate some of the kinematic changes observed in animals moving on different inclines (e.g. reduction in vertical oscillation of the center of mass and decreased stride length) underline the valuable and reasonable contributions that very simple, reductionist models, like the spring-mass model, can provide. [ABSTRACT FROM AUTHOR]

Locomotion in animals follows simple mathematical rules: when active, most time is spent at lower velocities and exponentially decreasing time is spent at increasing velocities; when inactive, there is an abundance of short quiescence periods decreasing by an inverse power function towards longer quiescence periods. We investigated whether these mathematical rules are 'universal', i.e. followed by all animals independently of their modes of movement (walking, gliding/crawling, swimming) or their habitat (land, water). Using automated video tracking with high temporal and spatial resolution, we investigated the velocity and quiescence period distributions in ten species representing major divisions of the animal kingdom (snails, woodlice, springtail, beetle, fish, reptile, chicken and mouse). We confirm that the two functions are followed by most species, although we also found important exceptions: the snails and woodlice showed uni- or bimodal velocity distributions; the snails did not follow the inverse power law of quiescence periods. Thus, most animals running on legs or swimming follow the expected distributions, while other runners (isopods) and substrate gliders/crawlers (snails) do not. We suggest a difference in velocity distribution between animals that use running to escape predators and those that do not (snails and woodlice). [ABSTRACT FROM AUTHOR]

Diving birds are regarded as a classic example of morphological convergence. Divers tend to have small wings extending from rotund bodies, requiring many volant species to fly with rapid wingbeats, and rendering others flightless. The high wing-loading of diving birds is frequently associated with the challenge of using forelimbs adapted for flight for locomotion in a "draggier" fluid, but this does not explain why species that rely exclusively on their feet to dive should have relatively small wings, as well. Therefore, others have hypothesized that ecological factors shared by wing-propelled and foot-propelled diving birds drive the evolution of high wing-loading. Following a reexamination of the aquatic habits of birds, we tested between hypotheses seeking to explain high wing-loading in divers using new comparative data and phylogenetically informed analyses. We found little evidence that wing-propelled diving selects for small wings, as wing-propelled and foot-propelled species share similar wing-loadings. Instead, our results suggest that selection to reduce buoyancy has driven high wing-loading in divers, offering insights for the development of bird-like aquatic robots. [ABSTRACT FROM AUTHOR]

Many animals frequently transition between different media while navigating their heterogeneous environments. These media vary in compliance, moisture content, and other characteristics that affect their physical properties. As a result, animals may need to alter their kinematics to adapt to potential changes in media while maintaining performance during predator escape and foraging. Due to its fluid nature, water is highly compliant, and although usually associated with swimming, water running has evolved in a variety of animals ranging from insects to mammals. While the best studied large water runners are the bipedal basilisk lizards (Basiliscus spp.), other lizards have also been observed to run across the surface of water, namely, Hemidactylus platyurus , a house gecko, and in this study, Anolis sagrei , the brown anole. Unlike the basilisk lizard, the primarily arboreal Anolis sagrei is not adapted for water running. Moreover, water running in A. sagrei , similar to that of the house gecko, was primarily quadrupedal. Here, we tested for performance and kinematic differences between aquatic and terrestrial running and if the variance in performance and kinematic variables differed between the two media. We found no difference in average and maximum velocity between running on land and water. We also found that Anolis sagrei had higher hindlimb stride frequencies, decreased duty factor, and shorter stride lengths on water, as well as more erect postures. Finally, we found that most kinematics did not differ in variance between the two media, but of those that were different, almost all were more variable during terrestrial running. Our findings show that animals may be capable of specialized modes of locomotion, even if they are not obviously adapted for them, and that they may do this by modulating their kinematics to facilitate locomotion through novel environments. [ABSTRACT FROM AUTHOR]

Kuznetsov and Panyutina (2022) offer a reanalysis of the kinematic and force plate data previously published by Bundle and Dial (2003). Their intention is to describe instantaneous wing forces during wing-assisted incline running (WAIR), focusing particularly on the upstroke phase. Based on their interpretation of wing forces and muscle function, the authors conclude that 'WAIR is a very specialized mode of locomotion that is employed by a few specialized birds as an adaptation to a very specific environment and involving highly developed flying features of the locomotor apparatus', and thus not relevant to the evolution of avian flight. Herein, we respond to the authors' interpretations, offering an alternative perspective on WAIR and, more generally, on studies exploring the evolution of avian flight. [ABSTRACT FROM AUTHOR]

Ecological physiologists and biomechanists have investigated swimming performance in a diversity of fishes; however, the connection between form, function, and energetics of locomotion has been rarely evaluated in the same system and under climate change scenarios. In this perspective, I argue that working within the framework of "EcoPhysioMechanics," i.e. integrating energetics and biomechanics tools, to measure locomotor performance and behavior under different abiotic factors, improves our understanding of the mechanisms, limits and costs of movement. To demonstrate how EcoPhysioMechanics can be applied to locomotor studies, I outline how linking biomechanics and physiology allows us to understand how fishes may modulate their movement to achieve high speeds or reduce the costs of locomotion. I also discuss how the framework is necessary to quantify swimming capacity under climate change scenarios. Finally, I discuss current dearth of integrative studies and gaps in empirical datasets that are necessary to understand fish swimming under changing environments. [ABSTRACT FROM AUTHOR]

Salamanders are often used as analogs for early tetrapods in paleontological reconstructions of locomotion. However, concerns have been raised about whether this comparison is justifiable, necessitating comparisons of a broader range of early tetrapods with salamanders. Here, we test whether the osteological morphology of the hindlimb in the early tetrapod (temnospondyl amphibian) Eryops megacephalus could have facilitated the sequence of limb configurations used by salamanders during terrestrial locomotion. To do so, we present a new method that enables the examination of full limb configurations rather than isolated joint poses. Based on this analysis, we conclude that E. megacephalus may indeed have been capable of salamander-like hindlimb kinematics. Our method facilitates the holistic visual comparison of limb configurations between taxa without reliance on the homology of coordinate system definitions, and can thus be applied to facilitate various comparisons between extinct and extant taxa, spanning the diversity of locomotion both past and present. [ABSTRACT FROM AUTHOR]

The capacity of certain animals to regrow a lost appendage has been exploited as a powerful tool to study development. As a result, we now understand many of the proximate details of the regeneration process. Ironically, despite being one of the oldest studied developmental phenomena, regeneration is not often considered in the context of natural selection and evolution. Why do select species retain the capacity to shed and regrow body parts, whereas more derived lineages do not? We conducted a comprehensive study on the costs and benefits of autotomy and regeneration on Hemigrapsus nudus , the purple shore crab. In the realms of feeding and locomotion, regeneration restored fitness to what it otherwise would have been; autotomized animals showed decreases in feeding and locomotion, but regenerated animals performed no differently than intact crabs. However, for fecundity and male–male competition, regenerated animals had the lowest fitness compared with control and autotomized crabs. Our results raise the intriguing possibility that tradeoffs associated with reproduction may have led to the loss of regenerative abilities in derived lineages such as mammals and birds. Future work on the hundreds of species that regenerate lost body parts will reveal if and how this hypothesis can address the pervasive speculation plaguing the ultimate causes of regenerative losses. [ABSTRACT FROM AUTHOR]

Early-life stress (ELS) can cause long-term effects on human health, ranging from adolescence to adulthood, and even to gerontic. Although clinical retrospective data suggest that ELS may be related to senile neurodegenerative diseases such as Parkinson's disease (PD), there are few prospective investigations to explore its real contribution to PD. Here, we investigated the behavioral, histochemical, neuromorphological, and transcriptional changes induced by maternal separation (MS), an ELS model. Without neurotoxin, MS rats showed behavioral alterations in olfaction, locomotion, and gait characters after depression compared with control rats. Based on neuroimaging and histochemistry, although we found that the dopaminergic system in the striatum was impaired after MS, the decrease of striatal dopamine level was ~33%. Consistently, tyrosine hydroxylase immunostaining positive neurons of MS rats in the substantia nigra showed deficit by about 20% in cell counting. Furthermore, using transcriptome sequencing, we discovered many differentially expressed genes (DEGs) of MS rats in the striatum significantly enriched in the pathway of dopaminergic synapse, and the biological process of locomotion and neuromuscular process controlling balance. Encouragingly, some representative DEGs relating to PD were singled out. These results suggest that ELS-depression rats potentially mimic some key features of prodromal stage of PD during natural senescence. In conclusion, our findings provide some novel insights into the future pathogenesis and therapeutic studies for PD related to depression. [ABSTRACT FROM AUTHOR]

Morphology is closely linked to locomotion and diet in animals. In animals that undertake long-distance migrations, limb morphology is under selection to maximize mobility and minimize energy expenditure. Migratory behaviours also interact with diet, such that migratory animals tend to be dietary generalists, whereas sedentary taxa tend to be dietary specialists. Despite a hypothesized link between migration status and morphology, phylogenetic comparative studies have yielded conflicting findings. We tested for evolutionary associations between migratory status and limb and bill morphology across kingbirds, a pan-American genus of birds with migratory, partially migratory and sedentary taxa. Migratory kingbirds had longer wings, in agreement with expectations that selection favours improved aerodynamics for long-distance migration. We also found an association between migratory status and bill shape, such that more migratory taxa had wider, deeper and shorter bills compared to sedentary taxa. However, there was no difference in intraspecific morphological variation among migrants, partial migrants and residents, suggesting that dietary specialization has evolved independently of migration strategy. The evolutionary links between migration, diet and morphology in kingbirds uncovered here further strengthen ecomorphological associations that underlie long-distance seasonal movements in animals. [ABSTRACT FROM AUTHOR]

We propose the use of bio-inspired robotics equipped with soft sensor technologies to gain a better understanding of the mechanics and control of animal movement. Soft robotic systems can be used to generate new hypotheses and uncover fundamental principles underlying animal locomotion and sensory capabilities, which could subsequently be validated using living organisms. Physical models increasingly include lateral body movements, notably back and tail bending, which are necessary for horizontal plane undulation in model systems ranging from fish to amphibians and reptiles. We present a comparative study of the use of physical modeling in conjunction with soft robotics and integrated soft and hyperelastic sensors to monitor local pressures, enabling local feedback control, and discuss issues related to understanding the mechanics and control of undulatory locomotion. A parallel approach combining live animal data with biorobotic physical modeling promises to be beneficial for gaining a better understanding of systems in motion. [ABSTRACT FROM AUTHOR]

Fishes overcome a variety of challenges in order to invade the terrestrial environment. Terrestrial invasions by fish occur over a variety of environmental contexts. In order to advance their bodies on land, fishes capable of terrestrial excursions tend to use one of three different types of locomotor modes: axial-based, appendage-based, or axial-appendage-based. Elongate species with reduced appendages, such as the American eel, Anguilla rostrata , rely on axial based locomotion in water and on land. When eels move from water to land as part of their complex life cycle, they inevitably encounter a variety of substrates and must traverse variable degrees of incline. The aim of this study was to determine the effect of substrate and incline on the terrestrial locomotion of the American eel. In order to do this, eels were filmed from a dorsal view on three substrates and four inclines: sand, loose pebbles, and fixed (glued) pebbles at 0°, 5°, 10°, and 15°. We digitized 20 evenly spaced points along the body to examine the following characteristics of locomotion: velocity, distance ratio (DR), and wave parameters such as wave amplitude, frequency, and length and assessed whether substrate, incline, or body position affected these parameters. DR, our metric of movement efficiency, was highest on the flat sand condition and lowest on 15° pebble conditions. Efficiency also varied across the body. Velocity followed a similar pattern being highest on sand at 0° and lowest at the steepest inclines. Wave amplitude generally increased toward the tail but was similar across substrates and inclines. Wave frequency was relatively consistent across the body on both pebble substrates, but on sand, frequency was higher toward the head but decreased toward the tail. Wavelengths on sand were the longest at 0° near the head and shorter wavelengths were observed on steeper inclines. Both pebble substrates elicited lower wavelengths that were more similar across the body. Overall, A. rostrata were more effective in navigating compliant substrates but struggled at steeper inclines. Our findings provide insight into locomotor challenges that American eels may encounter as they move from and between bodies of water. [ABSTRACT FROM AUTHOR]

Snakes have no limbs and can move in various environments using a simple elongated limbless body structure obtained through a long-term evolutionary process. Specifically, snakes have various locomotion patterns, which they change in response to conditions encountered. For example, on an unstructured terrain, snakes actively utilize the terrain's irregularities and move effectively by actively pushing their bodies against the "scaffolds" that they encounter. In a narrow aisle, snakes exhibit concertina locomotion, in which the tail part of the body is pulled forward with the head part anchored, and this is followed by the extension of the head part with the tail part anchored. Furthermore, snakes often exhibit three-dimensional (3-D) locomotion patterns wherein the points of ground contact change in a spatiotemporal manner, such as sidewinding and sinus-lifting locomotion. This ability is achieved possibly by a decentralized control mechanism, which is still mostly unknown. In this study, we address this aspect by employing a synthetic approach to understand locomotion mechanisms by developing mathematical models and robots. We propose a Tegotae -based decentralized control mechanism and use a 2-D snake-like robot to demonstrate that it can exhibit scaffold-based and concertina locomotion. Moreover, we extend the proposed mechanism to 3D and use a 3-D snake-like robot to demonstrate that it can exhibit sidewinding and sinus-lifting locomotion. We believe that our findings will form a basis for developing snake-like robots applicable to search-and-rescue operations as well as understanding the essential decentralized control mechanism underlying animal locomotion. [ABSTRACT FROM AUTHOR]

Small animals face a large challenge when running. A stable head is key to maintenance of a stable gaze and a good sense of self-motion and spatial awareness. However, trunk undulations caused by the cyclic limb movements result in involuntary head movements. Hence, the head needs to be stabilized. Humans are capable of stabilizing their head up to 2–3 Hz, but small animals run at cycle frequencies that are up to six times higher. We wondered how natural selection has adapted their head stabilization control. We observed that the relative contributions of vision, on the one hand, and vestibular perception and proprioception, on the other hand, remain the same when lizards undergo fast or slow body undulations in an experimental set-up. Lizards also maintain a short phase lag at both low and high undulation frequencies. Hence, we found no indication that they use a different control mechanism at high frequencies. Instead, head stabilization probably remains possible owing to faster reflex pathways and a lower head inertia. Hence, the intrinsic physical and neurological characteristics of lizards seem to be sufficient to enable head stabilization at high frequencies, obviating the need for evolutionary adaptation of the control pathways. These properties are not unique to lizards and might, therefore, also facilitate head stabilization at high frequencies in other small, fast animals. [ABSTRACT FROM AUTHOR]

Understanding how selective forces influence patterns of symmetry remains an active area of research in evolutionary biology. One hypothesis, which has received relatively little attention, suggests that the functional importance of morphological characters may influence patterns of symmetry. Specifically, it posits that for structures that display bilateral symmetry, those with greater functional importance should display lower levels of asymmetry. The aim of this study was to examine the patterns of fluctuating asymmetry (FA) present in the limb bones of freshwater turtles in the family Emydidae. Aquatic emydid turtles of the subfamily Deirochelyinae employ a hindlimb‐dominant swimming style, suggesting that hindlimbs should display lower levels of FA. Consistent with the morpho‐functional hypothesis of symmetry, we found a strong, clade‐wise pattern of humeral‐biased FA in aquatic Deirochelyinae. In contrast, some emydids of the subfamily Emydinae possess more terrestrial tendencies. As terrestrial locomotion places more equal importance on fore‐ and hindlimbs, we predicted that such behaviors may minimize differences in FA. No clade‐wise pattern was detected in the subfamily Emydinae. We also detected a phylogenetic signal in FA within the femur and discovered that FA has evolved at vastly different rates between the fore‐ and hindlimbs. [ABSTRACT FROM AUTHOR]

Environment and lifestyle induce substantial variation in the mechanisms of locomotion in vertebrates. A spectrum of adaptations related to locomotion is also present in limbless taxa, especially snakes, which have radiated successfully into a wide range of habitats. The majority of studies concerning habitat-driven variation in locomotor mechanisms of snakes have focused on the musculoskeletal system. Far less recognized is the variation in the morphology of ventral scales, which are another pivotal component of the locomotor system in snakes. Here, we investigated patterns of interspecific variation in the width of ventral scales in terms of lifestyle (hunting mode) and habitat occupied in 55 species of snakes belonging to eight families. We found that increasing terrestriality was associated with enlarged ventral scales. Reduction instead of maintenance of the width of ventral scales was observed in aquatic species, suggesting that wide ventral scales set constraints on aquatic locomotion. In terrestrial species, no significant differences were observed in terms of arboreality or hunting mode, which suggests overall optimization in the size of ventral scales towards terrestrial locomotion. Association between the width of ventral scales and locomotion can result in a habitat-dependent costs of abnormalities in ventral scale morphology, commonly observed in snakes. [ABSTRACT FROM AUTHOR]

Mammals flex, extend, and rotate their spines as they perform behaviors critical for survival, such as foraging, consuming prey, locomoting, and interacting with conspecifics or predators. The atlas–axis complex is a mammalian innovation that allows precise head movements during these behaviors. Although morphological variation in other vertebral regions has been linked to ecological differences in mammals, less is known about morphological specialization in the cervical vertebrae, which are developmentally constrained in number but highly variable in size and shape. Here, we present the first phylogenetic comparative study of the atlas–axis complex across mammals. We used spherical harmonics to quantify 3D shape variation of the atlas and axis across a diverse sample of species, and performed phylogenetic analyses to investigate if vertebral shape is associated with body size, locomotion, and diet. We found that differences in atlas and axis shape are partly explained by phylogeny, and that mammalian subclades differ in morphological disparity. Atlas and axis shape diversity is associated with differences in body size and locomotion; large terrestrial mammals have craniocaudally elongated vertebrae, whereas smaller mammals and aquatic mammals have more compressed vertebrae. These results provide a foundation for investigating functional hypotheses underlying the evolution of neck morphologies across mammals. [ABSTRACT FROM AUTHOR]

Patterns of phenotypic evolution can abruptly shift as species move between adaptive zones. Extant salamanders display three distinct life cycle strategies that range from aquatic to terrestrial (biphasic), to fully aquatic (paedomorphic) and to fully terrestrial (direct development). Life cycle variation is associated with changes in body form such as loss of digits, limb reduction or body elongation. However, the relationships among these traits and life cycle strategy remain unresolved. Here, we use a Bayesian modelling approach to test whether life cycle transitions by salamanders have influenced rates, optima and integration of primary locomotory structures (limbs and trunk). We show that paedomorphic salamanders have elevated rates of limb evolution with optima shifted towards smaller size and fewer digits compared to all other salamanders. Rate of hindlimb digit evolution is shown to decrease in a gradient as life cycles become more terrestrial. Paedomorphs have a higher correlation between hindlimb digit loss and increases in vertebral number, as well as reduced correlations between limb lengths. Our results support the idea that terrestrial plantigrade locomotion constrains limb evolution and, when lifted, leads to higher rates of trait diversification and shifts in optima and integration. The basic tetrapod body form of most salamanders and the independent losses of terrestrial life stages provide an important framework for understanding the evolutionary and developmental mechanisms behind major shifts in ecological zones as seen among early tetrapods during their transition from water to land. [ABSTRACT FROM AUTHOR]

Birds are a diverse and agile lineage of vertebrates that all use bipedal locomotion for at least part of their life. Thus birds provide a valuable opportunity to investigate how biomechanics and sensorimotor control are integrated for agile bipedal locomotion. This review summarizes recent work using terrain perturbations to reveal neuromechanical control strategies used by ground birds to achieve robust, stable, and agile running. Early experiments in running guinea fowl aimed to reveal the immediate intrinsic mechanical response to an unexpected drop ("pothole") in terrain. When navigating the pothole, guinea fowl experience large changes in leg posture in the perturbed step, which correlates strongly with leg loading and perturbation recovery. Analysis of simple theoretical models of running has further confirmed the crucial role of swing-leg trajectory control for regulating foot contact timing and leg loading in uneven terrain. Coupling between body and leg dynamics results in an inherent trade-off in swing leg retraction rate for fall avoidance versus injury avoidance. Fast leg retraction minimizes injury risk, but slow leg retraction minimizes fall risk. Subsequent experiments have investigated how birds optimize their control strategies depending on the type of perturbation (pothole, step, obstacle), visibility of terrain, and with ample practice negotiating terrain features. Birds use several control strategies consistently across terrain contexts: (1) independent control of leg angular cycling and leg length actuation, which facilitates dynamic stability through simple control mechanisms, (2) feedforward regulation of leg cycling rate, which tunes foot-contact timing to maintain consistent leg loading in uneven terrain (minimizing fall and injury risks), (3) load-dependent muscle actuation, which rapidly adjusts stance push-off and stabilizes body mechanical energy, and (4) multi-step recovery strategies that allow body dynamics to transiently vary while tightly regulating leg loading to minimize risks of fall and injury. In future work, it will be interesting to investigate the learning and adaptation processes that allow animals to adjust neuromechanical control mechanisms over short and long timescales. [ABSTRACT FROM AUTHOR]

The successful completion of many behaviors relies on sensory feedback. This symposium brought together researchers using novel techniques to study how different stimuli are encoded, how and where multimodal feedback is integrated, and how feedback modulates motor output in diverse modes of locomotion (aerial, aquatic, and terrestrial) in a diverse range of taxa (insects, fish, tetrapods), and in robots. Similar to biological organisms, robots can be equipped with integrated sensors and can rely on sensory feedback to adjust the output signal of a controller. Engineers often look to biology for inspiration on how animals have evolved solutions to problems similar to those experienced in robotic movement. Similarly, biologists too must proactively engage with engineers to apply computer and robotic models to test hypotheses and answer questions on the capacity and roles of sensory feedback in generating effective movement. Through a diverse group of researchers, including both biologists and engineers, the symposium attempted to catalyze new interdisciplinary collaborations and identify future research directions for the development of bioinspired sensory control systems, as well as the use of robots to test hypotheses in neuromechanics. [ABSTRACT FROM AUTHOR]

In many animals, movement often affects fitness components such as foraging or migration. On the other hand, individuals with higher mobility also often have fitness costs. This trade-off between movement and other traits may explain the maintenance of variation in moving ability in a population. However, few studies have focused on movement by walking, although many previous studies of insects have investigated the evolution of variations in mobility with wing polymorphism. In this study, we focused on the walking ability of the red flour beetle (Tribolium castaneum) and investigated whether females with higher than lower walking ability have fitness costs. The present results showed that females with genetically higher walking ability produced smaller eggs and had lower starvation resistance than females with lower walking ability. These results suggest that higher walking ability is costly for females, and this fitness cost may explain maintenance of variations of walking ability in a population. [ABSTRACT FROM AUTHOR]

Abstract: Fast‐growing genotypes living in time‐constrained environments are often more prone to predation, suggesting that growth‐predation risk trade‐offs are important factors maintaining variation in growth along climatic gradients. However, the mechanisms underlying how fast growth increases predation‐mediated mortality are not well understood. Here, we investigated if slow‐growing, low‐latitude individuals have faster escape swimming speed than fast‐growing high‐latitude individuals using common frog (Rana temporaria) tadpoles from eight populations collected along a 1500 km latitudinal gradient. We measured escape speed in terms of burst and endurance speeds in tadpoles raised in the laboratory at two food levels and in the presence and absence of a predator (Aeshna dragonfly larvae). We did not find any latitudinal trend in escape speed performance. In low food treatments, burst speed was higher in tadpoles reared with predators but did not differ between high‐food treatments. Endurance speed, on the contrary, was lower in high‐food tadpoles reared with predators and did not differ between treatments at low food levels. Tadpoles reared with predators showed inducible morphology (increased relative body size and tail depth), which had positive effects on speed endurance at low but not at high food levels. Burst speed was positively affected by tail length and tail muscle size in the absence of predators. Our results suggest that escape speed does not trade‐off with fast growth along the latitudinal gradient in R. temporaria tadpoles. Instead, escape speed is a plastic trait and strongly influenced by the interaction between resource level and predation risk. [ABSTRACT FROM AUTHOR]

Animals rarely move through their natural habitat without experiencing an environmental perturbation, such as an obstacle or drop in height. However, rapid perturbations due to mass changes are little studied, and these can have significant impacts on the ecology and evolution of animal locomotion due to associated changes in relative muscle mass, altered body shape, shifts in the position of the centre of mass and changes in the overall load on the animal. These ecologically important situations arise during gravidity, parturition/oviposition, feeding, load bearing and autotomy. How these ubiquitous changes shape the function of animals in their natural habitat will have profound impacts on survival, and are thus crucial for understanding the functional ecology of locomotion. We first discuss locomotor perturbations, with an emphasis on the duration associated with them. We then address how commonly studied modes of rapid mass changes (e.g. parturition, autotomy, feeding) perturb locomotion by examining the effects of natural and artificial changes in mass on locomotor performance, kinematics, kinetics and muscle function in terrestrial animals, with an emphasis on non-human vertebrates. After a detailed discussion of the functional ecology of rapid mass changes, we generate a framework for future studies, including predictions for why some animals are more robust to changes in mass. [ABSTRACT FROM AUTHOR]

Natural selection is thought to act on performance capacities and to shape morphology in a population, but which capacities are most ecologically relevant remains unclear. In nature, organisms encounter various substrates that incite changes in locomotor performance and thus natural selection may favour different morphological traits on different substrates, and may also shape substrate choice based on performance ability on each. We explored the relationships among morphology, performance and habitat use by examining maximal sprint performance and sprint sensitivity of six-lined racerunner (Aspidoscelis sexlineata) lizards on four substrates: rock, pebbles, sand and grass. We also identified morphological predictors of performance on each substrate. We found performance on grass, the most common microhabitat at the field site, to be suboptimal. Lizards tended to use substrates that optimized performance. Performance on different substrates was predicted by different morphological traits. Heterogeneity in available substrates on which individuals accomplish tasks important to fitness may result in selection on different morphological traits or selection to avoid suboptimal substrates compared to if only optimal substrates are used. [ABSTRACT FROM AUTHOR]

Insects are well equipped in walking on complex three-dimensional terrain, allowing them to overcome obstacles or catch prey. However, the gait transition for insects steering on a wall remains unexplored. Here, we find that honeybees adopted a tetrapod gait to change direction when climbing a wall. On the contrary to the common tripod gait, honeybees propel their body forward by synchronously stepping with both middle legs and then both front legs. This process ensures the angle of the central axis of the honeybee to be consistent with the crawling direction. Interestingly, when running in an alternating tripod gait, the central axis of honeybee sways around the center of mass under alternating tripod gait to maintain stability. Experimental results show that tripod, tetrapod, and random gaits result in the amazing consensus harmony on the climbing speed and gait stability, whether climbing on a smooth wall or walking on smooth ground. [ABSTRACT FROM AUTHOR]

Wild animals often engage in intense physical activity while performing tasks vital for their survival and reproduction associated with foraging, avoiding predators, fighting, providing parental care, and migrating. In this theme issue we consider how viewing these tasks as "exercise"--analogous to that performed by human athletes-may help provide insight into the mechanisms underlying individual variation in these types of behaviors and the importance of physical activity in an ecological context. In this article and throughout this issue, we focus on four key questions relevant to the study of behavioral ecology that may be addressed by studying wild animal behavior from the perspective of exercise physiology: (1) How hard do individual animals work in response to ecological (or evolutionary) demands?; (2) Do lab-based studies of activity provide good models for understanding activity in free-living animals and individual variation in traits?; (3) Can animals work too hard during "routine" activities?; and (4) Can paradigms of "exercise" and "training" be applied to free-living animals? Attempts to address these issues are currently being facilitated by rapid technological developments associated with physiological measurements and the remote tracking of wild animals, to provide mechanistic insights into the behavior of free-ranging animals at spatial and temporal scales that were previously impossible. We further suggest that viewing the behaviors of non-human animals in terms of the physical exercise performed will allow us to fully take advantage of these technological advances, draw from knowledge and conceptual frameworks already in use by human exercise physiologists, and identify key traits that constrain performance and generate variation in performance among individuals. It is our hope that, by highlighting mechanisms of behavior and performance, the articles in this issue will spur on further synergies between physiologists and ecologists, to take advantage of emerging cross-disciplinary perspectives and technologies. [ABSTRACT FROM AUTHOR]

Animals are constrained by their abilities and by interactions with environmental factors, such as low ambient temperatures. These constraints range from physical impossibilities to energetic inefficiencies, and may entail trade-offs. Some of the constraints related to locomotion and activity metabolism can be illustrated through allometric comparisons of mammals and lizards, as representative terrestrial vertebrate endotherms and ectotherms, respectively, because these lineages differ greatly in aerobic metabolic capacities, resting energetic costs, and thermoregulatory patterns. Allometric comparisons are both useful and unavoidable, but "outlier" species (unusual for their clade) can also inform evolutionary scenarios, as they help indicate extremes of possible adaptation within mammalian and saurian levels of organization. We compared mammals and lizards for standard metabolic rate (SMR), maximal oxygen consumption during forced exercise (VO2max), net (incremental) cost of transport (NCT), maximal aerobic speed (MAS), daily movement distance (DMD), daily energy expenditure (DEE) during the active season, and the ecological cost of transport (ECT=percentage of DEE attributable to locomotion). (Snakes were excluded because their limbless locomotion has no counterpart in terrestrial mammals.) We only considered lizard SMR, VO2max, NCT, MAS, and sprint speed data if measured at 35-40 °C. On average, MAS is ~7.4-fold higher in mammals, whereas SMR and VO2max are ~6-fold greater, but values for all three of these traits overlap (or almost overlap) between mammals and lizards, a fact that has not previously been appreciated. Previous studies show that sprint speeds are similar for smaller mammals and lizards, but at larger sizes lizards are not as fast as some mammals. Mammals move ~6-fold further each day than lizards, and DMD is by far the most variable trait considered here, but their NCT is similar. Mammals exceed lizards by ~11.4-fold for DEE. On average for both lineages, the ECT is surprisingly low, somewhat higher for lizards, and positively allometric. If a lizard and mammal of 100 g body mass were both to move their entire DMD at their MAS, they could do so in ~21 and 17 min, respectively, thus de-emphasizing the possible importance of time constraints. We conclude that ecological-energetic constraints related to locomotion are relatively more likely to occur in large, carnivorous lizards. Overall, our comparisons support the idea that the (gradual) evolution of mammalian endothermy did not necessarily require major changes in locomotor energetics, performance, or associated behaviors. Instead, we speculate that the evolution of thermoregulatory responses to low temperatures (e.g., shivering) may have been a key and "difficult" step in this transition. [ABSTRACT FROM AUTHOR]

Quantifying animal energy expenditure during locomotion in the field is generally based either on treadmill measurements or on estimates derived from a measured proxy. Two common proxies are heart rate (fH) and dynamic body acceleration (accelerometry). Both fH and accelerometry have been calibrated extensively under laboratory conditions, which typically involve prompting the animal to locomote on a treadmill at different speeds while simultaneously recording its rate of oxygen uptake (V̇O2) and the proxy. Field estimates of V̇O2 during locomotion obtained directly from treadmill running or from treadmill-calibrated proxies make assumptions about similarities between running in the field and in the laboratory. The present study investigated these assumptions, focusing on humans as a tractable species. First we investigated experimentally if and how the rate of energy expenditure during treadmill locomotion differs to that during field locomotion at the same speeds, with participants walking and running on a treadmill, on tarmac, and on grass, while wearing a mobile respirometry system. V̇O2 was substantially higher during locomotion in both of the field conditions compared with on a level treadmill: 9.1% on tarmac and 17.7% on grass. Second, we included these data in a meta-analysis of previous, related studies. The results were influenced by the studies excluded due to particulars of the experiment design, suggesting that participant age, the surface type, and the degree of turning during field locomotion may influence by how much treadmill and field locomotion V̇O2 differ. Third, based on our experiments described earlier, we investigated the accuracy of treadmill-calibrated accelerometry and fH for estimating V̇O2 in the field. The mean algebraic estimate errors varied between 10% and 35%, with the fH associated errors being larger than those derived from accelerometry. The mean algebraic errors were all underestimates of field V̇O2, by around 10% for fH and varying between 0% and 15% for accelerometry. Researchers should question and consider how accurately a treadmillderived proxy calibration of V̇O2 will estimate V̇O2 during terrestrial locomotion in free-living animals. [ABSTRACT FROM AUTHOR]

Studies of animal locomotion and movement largely assume that individuals are healthy and performing to the best of their abilities in ways which are adapted to their survival. However, wild animals face numerous ecological challenges that can compromise their health, reduce their performance capacity, impair their movement abilities and, ultimately, lower their fitness. By diverting resources and increasing host energetic demands, parasites, bacteria, and viruses (hereafter parasites) can dramatically influence the ways in which their hosts allocate energy to movement. Yet, the role of parasites in influencing animal locomotor performance and movement remains relatively unexplored, perhaps because animals often hide outward signs of sickness, and parasites tend to be small and inconspicuous to researchers. Here, we review how parasite infection can alter host locomotor performance via impacts on host morphology and physiology. We also give examples of behavioral strategies that some hosts employ to help overcome the disadvantages imposed by infection. Finally, we discuss how parasites can lead to both increased and decreased host movement patterns, either as an adaptive strategy for the host or due to manipulation by the parasite. The dynamic interplay between host movement (such as migration and dispersal) and infection has profound consequences for population and ecosystem-level processes that are influenced by movement. Acknowledging the important functional role played by parasites in driving the evolution of host locomotor performance and behavior is a critical step toward developing a comprehensive understanding of the causes and consequences of animal movement. [ABSTRACT FROM AUTHOR]

Wild animals maximize fitness through certain behaviors (e.g., foraging, mating, predator avoidance) that incur metabolic costs and often require high levels of locomotor activity. Consequently, the ability of animals to achieve high fitness often relies on their physiological capacity for exercise (aerobic scope) and/or their ability to acquire and utilize energy judiciously. Here, we explore how environmental factors and physiological limitations influence exercise and metabolism in fish while foraging, migrating to spawning grounds, and providing parental care. We do so with three case studies that use a number of approaches to studying exercise in wild fish using biologging and biotelemetry platforms. Bonefish (Albula vulpes) selectively use shallow water tropical marine environments to forage when temperatures are near optimal for aerobic scope and exercise capacity. Bonefish energy expenditure at upper thermal extremes is maximal while activity levels diminish, likely caused by reduced aerobic scope. Pacific salmon (Oncorhynchus spp.) reproductive migrations frequently involve passage through hydraulically challenging areas, and their ability to successfully pass these regions is constrained by their physiological capacity for exercise. Aerobic scope and swim performance are correlated with migration difficulty among sockeye salmon (O. nerka) populations; however, depletion of endogenous energy stores can also limit migration success. In another example, male smallmouth bass (Micropterus dolomieu) allocate a significant amount of energy to nest-guarding behaviors to protect their developing brood. Smallmouth bass body size, endogenous energy reserves, and physiological state influence nest-guarding behaviors and reproductive success. We suggest that in some scenarios (e.g., bonefish foraging, Pacific salmon dam passage) metabolic capacity for exercise may be the strongest determinant of biological fitness, while in others (e.g., long distance salmon migration, smallmouth bass parental care) energy stores may be more important. Interactions among environmental and ecological factors, fish behavior, and fish physiology offer important avenues of mechanistic inquiry to explain ecological dynamics and demonstrate how exercise is fundamental to the ecology of fish. [ABSTRACT FROM AUTHOR]

Despite strong purifying or directional selection, variation is ubiquitous in populations. One mechanism for the maintenance of variation is indirect genetic effects (IGEs), as the fitness of a given genotype will depend somewhat on the genes of its social partners. IGEs describe the effect of genes in social partners on the expression of the phenotype of a focal individual. Here, we ask what effect IGEs, and variation in IGEs between abiotic environments, has on locomotion in Drosophila. This trait is known to be subject to intralocus sexually antagonistic selection. We estimate the coefficient of interaction, Ψ, using six inbred lines of Drosophila. We found that Ψ varied between abiotic environments, and that it may vary across among male genotypes in an abiotic environment specific manner. We also found evidence that social effects of males alter the value of a sexually dimorphic trait in females, highlighting an interesting avenue for future research into sexual antagonism. We conclude that IGEs are an important component of social and sexual interactions and that they vary between individuals and abiotic environments in complex ways, with the potential to promote the maintenance of phenotypic variation. [ABSTRACT FROM AUTHOR]

Adjusting the efficiency of movement in response to environmental cues is an essential integrative characteristic of adaptive locomotion behavior across species. However, the modulatory molecules and the pathways involved are largely unknown. Recently, we demonstrated that in Caenorhabditis elegans, a loss-of-function of the two-pore-domain potassium (K2P) channel TWK-7 causes a fast, coordinated, and persistent forward crawling behavior in which five central aspects of stimulated locomotion--velocity, direction, wave parameters, duration, and straightness--are affected. Here, we isolated the reduction-of-function allele cau1 of the C. elegans gene kin-2 in a forward genetic screen and showed that it phenocopies the locomotor activity and locomotion behavior of twk-7(null) animals. Kin-2 encodes the negative regulatory subunit of protein kinase A (KIN-1/PKA). Consistently, we found that other gain-of-function mutants of the GαS-KIN-1/PKA pathway resemble kin-2(cau1) and twk-7(null) in locomotion phenotype. Using the powerful genetics of the C. elegans system in combination with cell type-specific approaches and detailed locomotion analyses, we identified TWK-7 as a putative downstream target of the GαS-KIN-1/PKA pathway at the level of the γ-aminobutyric acid (GABA)ergic D-type motor neurons. Due to this epistatic interaction, we suggest that KIN-1/PKA and TWK-7 may share a common pathway that is probably involved in the modulation of both locomotor activity and locomotion behavior during forward crawling. [ABSTRACT FROM AUTHOR]

Mesoplodont beaked whales are extreme divers, diving for over 45 mins and to depths of over 800 m. These dives are of similar depth and duration to those of the giant sperm whale (Physeter macrocephalus) whose body mass can be 50 times larger. Velten et al. (2013) provided anatomical data that demonstrated that on-board oxygen stores were sufficient to aerobically support the extreme dives of mesoplodonts if their diving metabolic rates are low. Because no physiological data yet exist, we utilized an anatomical approach—the body composition technique—to examine the relative metabolic rates of mesoplodonts. We utilized a systematic mass dissection protocol to compare the body composition of mesoplodonts with those of two short duration, shallow divers—the harbor porpoise (Phocoena phocoena) and bottlenose dolphin (Tursiops truncatus). We then investigated the body composition of two other extreme divers, the southern elephant seal (Mirounga leonina) and P. macrocephalus using data from the literature. Our results demonstrate that extreme divers invest a smaller percentage of their total body mass (TBM) in metabolically expensive brain and viscera, and a larger percent of their TBM in inexpensive integument, bone, and muscle, than do the shallow divers. Deep divers also share features of their locomotor muscle that contribute to relatively low tissue metabolic rates and high oxygen storage capacity, including large muscle fiber diameters, low mitochondrial volume densities, and high myoglobin concentrations. One feature of the locomotor muscle of mesoplodonts, though, is unique among deep divers investigated to date. Rather than having an endurance athlete's muscle fiber profile, dominated by slow oxidative fibers, mesoplodonts possess a sprinter's profile, dominated by fast glycolytic fibers. Velten et al. (2013) hypothesized that these fibers are likely inactive during routine swimming and provide a large, metabolically inexpensive oxygen store for the slow oxidative fibers to aerobically power swimming. We suggest that future anatomical analyses, coupled with performance data transduced through tagging studies, will enhance our understanding of the extreme diving capabilities of marine mammals. [ABSTRACT FROM AUTHOR]

The family Sciuridae is one of the most widespread and ecologically diverse lineages of rodents and represents an ideal model for investigating the evolution of locomotion modes and the historical biogeography of terrestrial mammals. We used a comprehensive database on locomotion modes, an updated phylogeny and novel biogeographic comparative methods to reassess the evolution of locomotion of squirrels and to investigate whether these locomotion modes evolved convergently in different continents. We found that locomotion changes occurred in different independent lineages of the family, including four reversals to terrestriality and one evolution of gliding. We also found evidence for Eurasia as the centre of origin of Sciuridae, challenging the classification of the oldest squirrel fossil records from the early Oligocene in North America. Additionally, Eurasia is also the possible centre of origin for most of squirrel subfamilies and tribes, and where locomotion changes have occurred. Parallel locomotion shifts could be explained by the adaptation towards different ecological niches followed by colonization of new continents. [ABSTRACT FROM AUTHOR]