Postural and respiratory activation of the trunk muscles changes with mode and speed of locomotion

Despite the importance of the deep intrinsic spinal muscles for trunk control, few studies have investigated their activity during human locomotion or how this may change with speed and mode of locomotion. Furthermore, it has not been determined whether the postural and respiratory functions, of which these muscles take part, can be coordinated when locomotor demands are increased. EMG recordings of abdominal and paraspinal muscles were made in seven healthy subjects using fine-wire and surface electrodes. Measurements were also made of respiration and gait parameters. Recordings were made for 10s as subjects walked on a treadmill at 1 and 2 ms(-1) and ran at 2, 3, 4 and 5 ms(-1). Unlike the superficial muscles, transversus abdominis was active tonically throughout the gait cycle with all tasks, except running at speeds of 3 ms(-1) and greater. All other muscles were recruited in a phasic manner. The relative duration of these bursts of activity was influenced by speed and/or mode of locomotion. Activity of all abdominal muscles, except rectus abdominis (RA), was modulated both for respiration and locomotor-related functions but this activity was affected by the speed and mode of locomotion. This study provides evidence that the deep abdominal muscles are controlled independently of the other trunk muscles. Furthermore, the pattern of recruitment of the trunk muscles and their respiratory and postural coordination is dependent on the speed and mode of locomotion. (C) 2003 Published by Elsevier B.V.

Locomotion and gaits of the northern brown bandicoot, Isoodon macrourus (Marsupalia: Peramelidae)

Gait repertoires of the northern brown bandicoot, Isoodon macrourus, were studied over a wide range of locomotor speeds. At low relative speeds, bandicoots used symmetrical gaits that included pacing, trotting, and lateral sequence strides. Forefoot contact duration was generally shorter than hind foot contact duration at all speeds. At moderate relative speeds bandicoots used half-bounding gaits with either no period of suspension or with a short gathered suspension. At high speeds the predominant gait had both a short extended and a short gathered suspension, although some strides comprised only an extended suspension. Increases in speed were accompanied by increases in spinal extension, presumably leading to the extended suspensions. On a stationary treadmill individuals occasionally used a bipedal gait. Maximum half-bounding speeds appear to be relatively low in this species.

Transient prenatal vitamin D deficiency is associated with hyperlocomotion in adult rats

Rat experiments have shown that prenatal Vitamin D deficiency leads to altered neonatal brain morphology, cell density and neurotrophin expression. In the current study we examined the hypothesis that Vitamin D deficiency during early development alters adult behaviour even when there is an intervening period in which the animal receives normal Vitamin D in later development. Rats were conceived and born to Vitamin D deficient dams (Birth); conceived, born and weaned from Vitamin D deficient dams (Weaning); or deficient in Vitamin D from conception to 10 weeks of age (Life). Litters were standardized to three males and three females per litter. All rat offspring were rendered normocalcaemic with calcium supplemented water (2 mM) after weaning. Control animals were born to mothers fed a normal diet but subject to similar litter size and calcium supplementation. At 10 weeks all animals were tested on the holeboard test, elevated plus maze test, social interaction observation, acoustic startle response test, prepulse inhibition of the acoustic startle response and a forced swim test. Early Vitamin D deficiency (Birth group) enhanced locomotion in the holeboard test and increased activity in the elevated plus maze. Thus, transient prenatal Vitamin D deficiency induces hyperlocomotion in adulthood, without severe motor abnormalities. (C) 2004 Elsevier B.V. All rights reserved.

Effect of prolonged inactivity on skeletal motor nerve terminals during aestivation in the burrowing frog, Cyclorana alboguttata

This study examined the effect of prolonged inactivity, associated with aestivation, on neuromuscular transmission in the green-striped burrowing frog, Cyclorana alboguttata. We compared the structure and function of the neuromuscular junctions on the iliofibularis muscle from active C. alboguttata and from C. alboguttata that had been aestivating for 6 months. Despite the prolonged period of immobility, there was no significant difference in the shape of the terminals (primary, secondary or tertiary branches) or the length of primary terminal branches between aestivators and non-aestivators. Furthermore, there was no significant difference in the membrane potentials of muscle fibres or in miniature end plate potential (EPP) frequency and amplitude. However, there was a significant decrease in evoked transmitter release characterised by a 56% decrease in mean EPP amplitude, and a 29% increase in the failure rate of nerve terminal action potentials to evoke transmitter release. The impact of this suite of neuromuscular characteristics on the locomotor performance of emergent frogs is discussed.

Interindividual Differences in Leg Muscle Mass and Pyruvate Kinase Activity Correlate with Interindividual Differences in Jumping Performance of Hyla multilineata

Frog jumping is an excellent model system for examining the structural basis of interindividual variation in burst locomotor performance. Some possible factors that affect jump performance, such as total body size, hindlimb length, muscle mass, and muscle mechanical and biochemical properties, were analysed at the interindividual (intraspecies) level in the tree frog Hyla multilineata. The aim of this study was to determine which of these physiological and anatomical variables both vary between individuals and are correlated with interindividual variation in jump performance. The model produced via stepwise linear regression analysis of absolute data suggested that 62% of the interindividual variation in maximum jump distance could be explained by a combination of interindividual variation in absolute plantaris muscle mass, total hindlimb muscle mass ( excluding plantaris muscle), and pyruvate kinase activity. When body length effects were removed, multiple regression indicated that the same independent variables explained 43% of the residual interindividual variation in jump distance. This suggests that individuals with relatively large jumping muscles and high pyruvate kinase activity for their body size achieved comparatively large maximal jump distances for their body size.

Changes in three dimensional lumbo-pelvic kinematics and trunk muscle activity with speed and mode of locomotion

Background Control of the trunk is critical for locomotor efficiency. However, investigations of trunk muscle activity and three-dimensional lumbo-pelvic kinematics during walking and running remain scarce. Methods. Gait parameters and three-dimensional lumbo-pelvic kinematics were recorded in seven subjects. Electromyography recordings of abdominal and paraspinal muscles were made using fine-wire and surface electrodes as subjects walked on a treadmill at 1 and 2 ms(-1) and ran at 2, 3, 4 and 5 ms(-1). Findings. Kinematic data indicate that the amplitude but not timing of lumbo-pelvic motion changes with locomotor speed. Conversely, a change in locomotor mode is associated with temporal but not spatial adaptation in neuromotor strategy. That is, peak transverse plane lumbo-pelvic rotation occurs at foot strike during walking but prior to foot strike during running. Despite this temporal change, there is a strong correlation between the amplitude of transverse plane lumbo-pelvic rotation and stride length during walking and running. In addition, Jumbo-pelvic motion was asymmetrical during all locomotor tasks. Trunk muscle electromyography occurred biphasically in association with foot strike. Transversus abdominis was tonically active with biphasic modulation. Consistent with the kinematic data, electromyography activity of the abdominal muscles and the superficial fibres of multifidus increased with locomotor speed, and timing of peak activity of superficial multifidus and obliquus externus abdominis was modified in association with the temporal adaptation in lumbo-pelvic motion with changes in locomotor mode. Interpretation. These data provide evidence of the association between lumbo-pelvic motion and trunk muscle activity during locomotion at different speeds and modes. (c) 2005 Elsevier Ltd. All rights reserved.

Behavioural characterization of Vitamin D receptor knockout mice

Vitamin D (calcitriol) is a nuclear transcription regulator acting via a nuclear hormone receptor (VDR). In addition to its role in the regulation of calcium and phosphate horneostasis and in bone formation, Vitamin D is also thought to be involved in brain function. The aim of this study was to behaviourally phenotype VDR knockout mice. We characterized the behaviour of VDR null mutant mice and wildtype littermate controls by subjecting them to a range of tests including a primary behavioural screen (using the SHIRPA protocol), rotarod, gait analysis, Y-maze, marble burying test, bedding test, holeboard test, elevated plus maze, open field test and prepulse inhibition of the acoustic startle response. There were no effects of genotype on most of the scores from the SHIRPA protocol except that VDR -/- mice had alopecia, were shorter and weighed less than VDR +/+ mice. VDR -/- mice had a shorter gait as well as impairments on the rotarod, in the bedding test and impaired habituation in both the open field and on the acoustic startle response. The VDR -/- mice had normal acoustic startle responses but had impaired PPI at long (256 ms) but not short (64 ms) prepulse to pulse intervals. The VDR -/- mice were less active in the open field and buried fewer marbles in the marble burying test. However, there were no differences in the time spent on the open arms of the elevated plus maze or in working memory as assessed by repeat arm entries on the Y-maze. Therefore, it appears that VDR -/- mice have muscular and motor impairments that significantly affects locomotor behaviour but seemingly no impairments in cognition as indicated by exploration, working memory or anxiety. (C) 2004 Elsevier B.V. All rights reserved.

Post-hatching growth and development of the pectoral and pelvic limbs in the black noddy, Anous minutus

The black tern (Anous minutus) uses a semi-precocial growth strategy. Terrestrial locomotor capacity occurs soon after hatching, but pectoral limb development is delayed and flight is not possible until about post-hatching day 50. A growth series (hatchlings to fledglings) was used to explore how limb musculoskeletal development varied with body mass. In the pelvic limb, bone lengths scaled isometrically or with negative allometry. Gastrocnemius muscle mass and the failure load and stiffness of the tibiotarsus scaled isometrically. In the pectoral limb, pectoralis and supracoracoideus muscle masses increased with strong positive allometry that was mirrored by increases in wing bone strength and stiffness. Bending strength (σult) and modulus (E) remained fairly constant throughout development to fledging for all limb bones. The moment of inertia (I) scaled with negative allometry for the tibiotarsus and with strong positive allometry in the wing bones. Differences in σult and E of the tibiotarsus between pre-fledged chicks and adults was due, primarily, to increases in bone density rather than increases in the moment of inertia of the skeletal elements, whereas σult of wing bones was a function of increases in both bone density and I. Early development of functional pelvic limbs in tree-nesting birds is relatively unusual, and presumably reflects a familial trait that does not appear to compromise breeding success in this species.

Coadaptation: A unifying principle in evolutionary thermal biology

Over the last 50 yr, thermal biology has shifted from a largely physiological science to a more integrated science of behavior, physiology, ecology, and evolution. Today, the mechanisms that underlie responses to environmental temperature are being scrutinized at levels ranging from genes to organisms. From these investigations, a theory of thermal adaptation has emerged that describes the evolution of thermoregulation, thermal sensitivity, and thermal acclimation. We review and integrate current models to form a conceptual model of coadaptation. We argue that major advances will require a quantitative theory of coadaptation that predicts which strategies should evolve in specific thermal environments. Simply combining current models, however, is insufficient to understand the responses of organisms to thermal heterogeneity; a theory of coadaptation must also consider the biotic interactions that influence the net benefits of behavioral and physiological strategies. Such a theory will be challenging to develop because each organism's perception of and response to thermal heterogeneity depends on its size, mobility, and life span. Despite the challenges facing thermal biologists, we have never been more pressed to explain the diversity of strategies that organisms use to cope with thermal heterogeneity and to predict the consequences of thermal change for the diversity of communities.

Effect of thermal acclimation on female resistance to forced matings in the eastern mosquitofish

All copulations in the eastern mosquitofish, Gambusia holbrooki, are coercive-and-achieved by force. Female G. holbrooki never appear to cooperate with males, but vigorously resist matings at all times. We examined the role of females within a sexually coercive mating system by investigating the ability of female G. holbrooki to resist forced copulations after acclimation to 16 degrees C and 32 degrees C for 4-5 weeks. We also examined burst swimming performance of female G. holbrooki after acclimation, as this trait is likely to underlie a female's ability to resist forced matings. We predicted that if female G. holbrooki indiscriminately resist matings from all males, acclimation would enhance female resistance at their acclimation temperature. However, we found that it did not. We also predicted that if females are able to influence the outcome of mating interactions, acclimation to an optimal thermal environment may induce females to reduce resistance. In support of this prediction, females acclimated at 32 degrees C were able to modify their resistance behaviour between exposure to 16 degrees C and 32 degrees C. The rate of copulations experienced by 32 inverted perpendicular C-acclimated females was 2.5 times greater at 32 degrees C than at 16 degrees C. In addition, acclimation at 32 degrees C significantly enhanced burst swimming performance at 32 degrees C but no effect of acclimation was observed at 16 degrees C. Our results suggest that female G. holbrooki are able to play a greater role in determining the outcome of sexual coercive mating interactions than previously thought. (c) 2006 The Association for the Shidy of Animal Behavioor. Published by Elsevier Ltd. All rights reserved.

Influence of incubation temperature on hatchling phenotype in reptiles

Incubation temperature influences hatchling phenotypes such as sex, size, shape, color, behavior, and locomotor performance in many reptiles, and there is growing concern that global warming might adversely affect reptile populations by altering frequencies of hatchling phenotypes. Here I overview a recent theoretical model used to predict hatchling sex of reptiles with temperature-dependent sex determination. This model predicts that sex ratios will be fairly robust to moderate global warming as long as eggs experience substantial daily cyclic fluctuations in incubation temperatures so that embryos are exposed to temperatures that inhibit embryonic development for part of the day. I also review studies that examine the influence of incubation temperature on posthatch locomotion performance and growth because these are the traits that are likely to have the greatest effect on hatchling fitness. The majority of these studies used artificial constant-temperature incubation, but some have addressed fluctuating incubation temperature regimes. Although the number of studies is small, it appears that fluctuating temperatures may enhance hatchling locomotor performance. This finding should not be surprising, given that the majority of natural reptile nests are relatively shallow and therefore experience daily fluctuations in incubation temperature.

On the tracks of the earliest dinosaurs: implications for the hypothesis of dinosaurian monophyly

From the record of dinosaurian skeletal remains it has been inferred that the origin and initial diversification of dinosaurs were rapid events, occupying an interval of about 5 million years in the Late Triassic. By contrast numerous reports of dinosauroid tracks imply that the emergence of dinosaurs was a more protracted affair extending through much of the Early and Middle Triassic. This study finds no convincing evidence of dinosaur tracks before the late Ladinian. Each of the three dinosaurian clades - Theropoda, Sauropodomorpha, Ornithischia - produced a unique track morphotype that appears to be an independent modification of the chirotherioid pattern attributed to stem-group archosaurs (thecodontian reptiles). The existence of three divergent track morphotypes is consistent with the concept of dinosaurian polyphyly but can be reconciled with the hypothesis of dinosaurian monophyly only by invoking many and rapid reversals in the locomotor anatomy of early dinosaurs. The origin of dinosaurs was not the correlate or consequence of any single event or process, be it global change, competitive replacement, or opportunism in the wake of mass extinction. Instead the origin of dinosaurs is envisaged as a series of three cladogenetic events over an interval of at least 10 million years and possibly as much as 25 million years. This scenario of dinosaurian polyphyly is as well-supported by fossil evidence as is the currently favoured view of dinosaurian monophyly.

Short- and long-term consequences of thermal variation in the larval environment of anurans

To survive adverse or unpredictable conditions in the ontogenetic environment, many organisms retain a level of phenotypic plasticity that allows them to meet the challenges of rapidly changing conditions. Larval anurans are widely known for their ability to modify behaviour, morphology and physiological processes during development, making them an ideal model system for studies of environmental effects on phenotypic traits. Although temperature is one of the most important factors influencing the growth, development and metamorphic condition of larval anurans, many studies have failed to include ecologically relevant thermal fluctuations among their treatments. We compared the growth and age at metamorphosis of striped marsh frogs Limnodynastes peronii raised in a diurnally fluctuating thermal regime and a stable regime of the same mean temperature. We then assessed the long-term effects of the larval environment on the morphology and performance of post-metamorphic frogs. Larval L. peronii from the fluctuating treatment were significantly longer throughout development and metamorphosed about 5 days earlier. Frogs from the fluctuating group metamorphosed at a smaller mass and in poorer condition compared with the stable group, and had proportionally shorter legs. Frogs from the fluctuating group showed greater jumping performance at metamorphosis and less degradation in performance during a 10-week dormancy. Treatment differences in performance could not be explained by whole-animal morphological variation, suggesting improved contractile properties of the muscles in the fluctuating group.

Swimming behaviour and post-swimming activity in Vitamin D receptor knockout mice

Animal experiments have shown that Vitamin D plays a role in both brain development and adult brain function. The adult Vitamin D receptor null mutant mouse (VDR -/-) is reported to be less active and more anxious than wild-type litter mate controls and to have poor swimming ability. However, an anxious behavioural phenotype is inferred from differences in locomotor behaviour. This is a general problem in behavioural phenotyping where a neurological phenotype is inferred from changes in locomotion which will be affected by non-neurological factors, such as muscle fatigue. In this study of VDR -/-, we conducted a detailed examination of one form of motor behaviour, swimming, compared to wildtype littermate controls. Swimming was assessed using a forced swim test, a laneway swimming test and a watermaze test using a visible platform. Post-swimming activity was assessed by comparing grooming and rearing behaviour before, and 5 min after, the forced swimming test. We replicated previous findings in which VDR -/- mice demonstrate more sinking episodes than wildtype controls in the forced swim test but they were similar to controls in the time taken to swim a 1 m laneway, and in the time taken to reach a visible platform in the watermaze. Thus, the VDR -/- mice were able to swim but were not able to float. Grooming and rearing behaviour of the VDR -/- mice was similar to wildtype controls before the forced swim but the VDR -/- were much less active after the swim compared with wildtype mice which displayed high levels of grooming and rearing. We conclude that VDR -/- mice have muscular and motor impairments that do not affect their ability to swim but significantly alters the ability to float as well as their post-swimming activity. Differences in muscle strength may confound tests of activity that are used to infer an anxious phenotype. (c) 2005 Elsevier Inc. All rights reserved.

Swimming performance of hatchling green turtles is affected by incubation temperature

In an experiment repeated for two separate years, incubation temperature was found to affect the body size and swimming performance of hatchling green turtles (Chelonia mydas). In the first year, hatchlings from eggs incubated at 26 degrees C were larger in size than hatchlings from 28 and 30 degrees C, whilst in the second year hatchlings from 25.5 degrees C were similar in size to hatchings from 30 degrees C. Clutch of origin influenced the size of hatchlings at all incubation temperatures even when differences in egg size were taken into account. In laboratory measurements of swimming performance, in seawater at 28 degrees C, hatchlings from eggs incubated at 25.5 and 26 degrees C had a lower stroke rate frequency and lower force output than hatchlings from 28 and 30 degrees C. These differences appeared to be caused by the muscles of hatchlings from cooler temperatures fatiguing at a faster rate. Clutch of origin did not influence swimming performance. This finding that hatchling males incubated at lower temperature had reduced swimming ability may affect their survival whilst running the gauntlet of predators in shallow near-shore waters, prior to reaching the relative safety of the open sea.