Integration of biomechanical compliance, leverage, and power in elephant limbs.
ABSTRACT: The structure and motion of elephant limbs are unusual compared with those of other animals. Elephants stand and move with straighter limbs (at least when walking), and have limited speed and gait. We devised novel experiments to examine how the limbs of elephants support and propel their mass and to explore the factors that may constrain locomotor performance in these largest of living land animals. We demonstrate that elephant limbs are remarkably compliant even in walking, which maintains low peak forces. Dogma defines elephant limbs as extremely "columnar" for effective weight support, but we demonstrate that limb effective mechanical advantage (EMA) is roughly one-third of that predicted for their size. EMA in elephants is actually smaller than that in horses, which are only one-tenth their mass; it is comparable to human limb values. EMA drops sharply with speed in elephants, as it does in humans. Muscle forces therefore must increase as the limbs become more flexed, and we show how this flexion translates to greater volumes of muscle recruited for locomotion and hence metabolic cost. Surprisingly, elephants use their forelimbs and hindlimbs in similar braking and propulsive roles, not dividing these functions among limbs as was previously assumed or as in other quadrupeds. Thus, their limb function is analogous to four-wheel-drive vehicles. To achieve the observed limb compliance and low peak forces, elephants synchronize their limb dynamics in the vertical direction, but incur considerable mechanical costs from limbs working against each other horizontally.
Project description:Synopsis Animal legs are diverse, complex, and perform many roles. One defining requirement of legs is to facilitate terrestrial travel with some degree of economy. This could, theoretically, be achieved without loss of mechanical energy if the body could take a continuous horizontal path supported by vertical forces only—effectively a wheel-like translation, and a condition closely approximated by walking tortoises. If this is a potential strategy for zero mechanical work cost among quadrupeds, how might the structure, posture, and diversity of both sprawled and parasagittal legs be interpreted? In order to approach this question, various linkages described during the industrial revolution are considered. Watt’s linkage provides an analogue for sprawled vertebrates that uses diagonal limb support and shows how vertical-axis joints could enable approximately straight-line horizontal translation while demanding minimal mechanical power. An additional vertical-axis joint per leg results in the wall-mounted pull-out monitor arm and would enable translation with zero mechanical work due to weight support, without tipping or toppling. This is consistent with force profiles observed in tortoises. The Peaucellier linkage demonstrates that parasagittal limbs with lateral-axis joints could also achieve the zero-work strategy. Suitably tuned four-bar linkages indicate this is feasibly approximated for flexed, biologically realistic limbs. Where “walking” gaits typically show out of phase fluctuation in center of mass kinetic and gravitational potential energy, and running, hopping or trotting gaits are characterized by in-phase energy fluctuations, the zero limb-work strategy approximated by tortoises would show zero fluctuations in kinetic or potential energy. This highlights that some gaits, perhaps particularly those of animals with sprawled or crouched limbs, do not fit current kinetic gait definitions; an additional gait paradigm, the “zero limb-work strategy” is proposed.
Project description:Joint surfaces of limb bones are loaded in compression by reaction forces generated from body weight and musculotendon complexes bridging them. In general, joints of eutherian mammals have regions of high radiodensity subchondral bone that are better at resisting compressive forces than low radiodensity subchondral bone. Identifying similar form-function relationships between subchondral radiodensity distribution and joint load distribution within the marsupial postcranium, in addition to providing a richer understanding of marsupial functional morphology, can serve as a phylogenetic control in evaluating analogous relationships within eutherian mammals. Where commonalities are established across phylogenetic borders, unifying principles in mammalian physiology, morphology, and behavior can be identified. Here, we assess subchondral radiodensity patterns in distal tibiae of several marsupial taxa characterized by different habitual activities (e.g., locomotion). Computed tomography scanning, maximum intensity projection maps, and pixel counting were used to quantify radiodensity in 41 distal tibiae of bipedal (5 species), arboreal quadrupedal (4 species), and terrestrial quadrupedal (5 species) marsupials. Bipeds (Macropus and Wallabia) exhibit more expansive areas of high radiodensity in the distal tibia than arboreal (Dendrolagus, Phascolarctos, and Trichosurus) or terrestrial quadrupeds (Sarcophilus, Thylacinus, Lasiorhinus, and Vombatus), which may reflect the former carrying body weight only through the hind limbs. Arboreal quadrupeds exhibit smallest areas of high radiodensity, though they differ non-significantly from terrestrial quadrupeds. This could indicate slightly more compliant gaits by arboreal quadrupeds compared to terrestrial quadrupeds. The observed radiodensity patterns in marsupial tibiae, though their statistical differences disappear when controlling for phylogeny, corroborate previously documented patterns in primates and xenarthrans, potentially reflecting inferred limb use during habitual activities such as locomotion. Despite the complex nature of factors contributing to joint loads, broad observance of these patterns across joints and across a variety of taxa suggests that subchondral radiodensity can be used as a unifying form-function principle within Mammalia.
Project description:Background:Elephant camps are among the most attractive destinations in Thailand for tourists from many countries. A wide range of management strategies are used by these camps, which can have varied impacts on health and welfare of elephants. Methods:This study surveyed 33 camps with 627 elephants in northern Thailand to quantify the types of management practices and work activities experienced by captive elephants. The survey consisted of an interview with camp owners, and direct observations of camp operations. Results:Data revealed considerable variation in elephant demographics, work activities, elephant care (i.e., housing, restraint, nutrition, health care, and breeding), and mahout management among the camps. In general, older camps (those in existence for >16 years) were involved in more intensive activities, like riding with saddles and shows. By contrast, newer camps provided more one-on-one activities for tourists and elephants, and emphasized more intimate, relaxing experiences (e.g., feeding, bathing, walking) than entertainment. A demographic shift also was observed, with elephants 20 years of age and younger having a sex ratio closer to 1:1 compared to elephants in older age categories (1:4.1-1:9.8). Discussion:Shifts in elephant management to less intensive activities were observed, which could have positive implications for elephant welfare. The shifting sex ratio suggests successful captive breeding is resulting in the birth of more males, which could present new welfare challenges in the future, because bulls can be more difficult to manage and socialize, and are more likely to be kept isolated during musth. Ultimately, the goal is to understand how camp activities affect welfare, and to develop science-based guidelines and standards to aid in the management of both male and female elephants used in tourism.
Project description:Limb movements are generally driven by active muscular contractions working with and against passive forces arising in muscles and other structures. In relatively heavy limbs, the effects of gravity and inertia predominate, whereas in lighter limbs, passive forces intrinsic to the limb are of greater consequence. The roles of passive forces generated by muscles and tendons are well understood, but there has been little recognition that forces originating within joints themselves may also be important, and less still that these joint forces may be adapted through evolution to complement active muscle forces acting at the same joint.We examined the roles of passive joint forces in insect legs with different arrangements of antagonist muscles. We first show that passive forces modify actively generated movements of a joint across its working range, and that they can be sufficiently strong to generate completely passive movements that are faster than active movements observed in natural behaviors. We further demonstrate that some of these forces originate within the joint itself. In legs of different species adapted to different uses (walking, jumping), these passive joint forces complement the balance of strength of the antagonist muscles acting on the joint. We show that passive joint forces are stronger where they assist the weaker of two antagonist muscles.In limbs where the dictates of a key behavior produce asymmetry in muscle forces, passive joint forces can be coadapted to provide the balance needed for the effective generation of other behaviors.
Project description:Walking is a recommended form of physical activity for obese adults, yet the effects of obesity and walking speed on the biomechanics of walking are not well understood. The purpose of this study was to examine joint kinematics, muscle force requirements and individual muscle contributions to the walking ground reaction forces (GRFs) at two speeds (1.25 ms(-1) and 1.50 ms(-1)) in obese and nonobese adults. Vasti (VAS), gluteus medius (GMED), gastrocnemius (GAST), and soleus (SOL) forces and their contributions to the GRFs were estimated using three-dimensional musculoskeletal models scaled to the anthropometrics of nine obese (35.0 (3.78 kg m(-2))); body mass index mean (SD)) and 10 nonobese (22.1 (1.02 kg m(-2))) subjects. The obese individuals walked with a straighter knee in early stance at the faster speed and greater pelvic obliquity during single limb support at both speeds. Absolute force requirements were generally greater in obese vs. nonobese adults, the main exception being VAS, which was similar between groups. At both speeds, lean mass (LM) normalized force output for GMED was greater in the obese group. Obese individuals appear to adopt a gait pattern that reduces VAS force output, especially at speeds greater than their preferred walking velocity. Greater relative GMED force requirements in obese individuals may contribute to altered kinematics and increased risk of musculoskeletal injury/pathology. Our results suggest that obese individuals may have relative weakness of the VAS and hip abductor muscles, specifically GMED, which may act to increase their risk of musculoskeletal injury/pathology during walking, and therefore may benefit from targeted muscle strengthening.
Project description:Elephant camps are among the most popular destinations in Thailand for tourists from many countries. A wide range of management strategies are used by these camps, which can have varied impacts on health and welfare of elephants. The objectives of this study were to examine relationships between FGM (fecal glucocorticoid metabolite) concentrations and camp management factors (work routine, walking, restraint, rest area, foraging), and to other welfare indicators (stereotypic behaviors, body condition, foot health, and skin wounds). Data were obtained on 84 elephants (18 males and 66 females) from 15 elephant camps over a 1-year period. Elephants were examined every 3 months and assigned a body condition score, foot score, and wound score. Fecal samples were collected twice monthly for FGM analysis. Contrary to some beliefs, elephants in the observation only program where mahouts did not carry an ankus for protection had higher FGM concentrations compared to those at camps that offered riding with a saddle and shows. Elephants that were tethered in the forest at night had lower FGM concentrations compared to elephants that were kept in open areas inside the camps. There was an inverse relationship between FGM concentrations and occurrence of stereotypy, which was not anticipated. Thus, assessing adrenal activity via monitoring of FGM concentrations can provide important information on factors affecting the well-being of elephants. Results suggest that more naturalistic housing conditions and providing opportunities to exercise may be good for elephants under human care in Thailand, and that a no riding, no hook policy does not necessarily guarantee good welfare.
Project description:Most of southern Africa's elephants inhabit environments where environmental temperatures exceed body temperature, but we do not know how elephants respond to such environments. We evaluated the relationships between apparent thermoregulatory behaviour and environmental, skin and core temperatures for tame savanna elephants (Loxodonta africana) that were free-ranging in the hot parts of the day, in their natural environment. Environmental temperature dictated elephant behaviour within a day, with potential consequences for fine-scale habitat selection, space use and foraging. At black globe temperatures of ~30°C, elephants adjusted their behaviour to reduce environmental heat load and increase heat dissipation (e.g. shade use, wetting behaviour). Resting, walking and feeding were also influenced by environmental temperature. By relying on behavioural and autonomic adjustments, the elephants maintained homeothermy, even at environmental temperatures exceeding 40°C. Elephants clearly have the capacity to deal with extreme heat, at least in environments with adequate resources of forage, water and shade. Future conservation actions should provide for the thermoregulatory, resource and spatial needs of elephants.
Project description:BACKGROUND: Previous phylogenetic analyses of African elephants have included limited numbers of forest elephant samples. A large-scale assessment of mitochondrial DNA diversity in forest elephant populations here reveals a more complex evolutionary history in African elephants as a whole than two-taxon models assume. RESULTS: We analysed hypervariable region 1 of the mitochondrial control region for 71 new central African forest elephants and the mitochondrial cytochrome b gene from 28 new samples and compare these sequences to other African elephant data. We find that central African forest elephant populations fall into at least two lineages and that west African elephants (both forest and savannah) share their mitochondrial history almost exclusively with central African forest elephants. We also find that central African forest populations show lower genetic diversity than those in savannahs, and infer a recent population expansion. CONCLUSION: Our data do not support the separation of African elephants into two evolutionary lineages. The demographic history of African elephants seems more complex, with a combination of multiple refugial mitochondrial lineages and recurrent hybridization among them rendering a simple forest/savannah elephant split inapplicable to modern African elephant populations.
Project description:With populations of African savannah elephants (Loxodonta africana) declining across the continent, assessing the status of individual elephant populations is important for conservation. Angola's elephant population represents a key linkage between the larger populations of Namibia and Botswana. Elephants in Angola were decimated during the 1975-2002 Angolan civil war, but a 2005 survey showed that populations were recolonizing former habitats. Between 2005 and 2015, no research was permitted on elephants in Angola, but elsewhere in Africa many elephant populations experienced a poaching crisis. In 2015, we were able to resume elephant research in Angola. We used aerial surveys and satellite monitoring of collared elephants to determine the current status of elephant populations in Angola and to learn how human populations may be affecting elephant habitat usage. The aerial survey revealed a population of 3,395 ± SE of 797 elephants, but populations had declined 21% from the 2005 estimate. The high number of carcasses observed on the survey suggests that populations may have increased after the 2005 survey but were declining rapidly as of 2015. Satellite-collared elephants avoided areas <6 km from human indicators but preferred areas nearer humans at scales of 6-40 km, suggesting that humans may be displacing elephants from preferred habitats near rivers. Taken together, these results suggest that Angola's elephant population is experiencing intense poaching and may be losing habitat to human settlements. Without action to conserve their populations, Angola's elephants face an uncertain future.
Project description:African elephants (Loxodonta africana) are imperiled by poaching and habitat loss. Despite global attention to the plight of elephants, their population sizes and trends are uncertain or unknown over much of Africa. To conserve this iconic species, conservationists need timely, accurate data on elephant populations. Here, we report the results of the Great Elephant Census (GEC), the first continent-wide, standardized survey of African savannah elephants. We also provide the first quantitative model of elephant population trends across Africa. We estimated a population of 352,271 savannah elephants on study sites in 18 countries, representing approximately 93% of all savannah elephants in those countries. Elephant populations in survey areas with historical data decreased by an estimated 144,000 from 2007 to 2014, and populations are currently shrinking by 8% per year continent-wide, primarily due to poaching. Though 84% of elephants occurred in protected areas, many protected areas had carcass ratios that indicated high levels of elephant mortality. Results of the GEC show the necessity of action to end the African elephants' downward trajectory by preventing poaching and protecting habitat.