Showing papers in "The Journal of Experimental Biology in 2006"

Phenotypic plasticity and evolution by genetic assimilation

Abstract: In addition to considerable debate in the recent evolutionary literature about the limits of the Modern Synthesis of the 1930s and 1940s, there has also been theoretical and empirical interest in a variety of new and not so new concepts such as phenotypic plasticity, genetic assimilation and phenotypic accommodation. Here we consider examples of the arguments and counter-arguments that have shaped this discussion. We suggest that much of the controversy hinges on several misunderstandings, including unwarranted fears of a general attempt at overthrowing the Modern Synthesis paradigm, and some fundamental conceptual confusion about the proper roles of phenotypic plasticity and natural selection within evolutionary theory.

Frictional adhesion: A new angle on gecko attachment.

Abstract: Directional arrays of branched microscopic setae constitute a dry adhesive on the toes of pad-bearing geckos, nature's supreme climbers. Geckos are easily and rapidly able to detach their toes as they climb. There are two known mechanisms of detachment: (1) on the microscale, the seta detaches when the shaft reaches a critical angle with the substrate, and (2) on the macroscale, geckos hyperextend their toes, apparently peeling like tape. This raises the question of how geckos prevent detachment while inverted on the ceiling, where body weight should cause toes to peel and setal angles to increase. Geckos use opposing feet and toes while inverted, possibly to maintain shear forces that prevent detachment of setae or peeling of toes. If detachment occurs by macroscale peeling of toes, the peel angle should monotonically decrease with applied force. In contrast, if adhesive force is limited by microscale detachment of setae at a critical angle, the toe detachment angle should be independent of applied force. We tested the hypothesis that adhesion is increased by shear force in isolated setal arrays and live gecko toes. We also tested the corollary hypotheses that (1) adhesion in toes and arrays is limited as on the microscale by a critical angle, or (2) on the macroscale by adhesive strength as predicted for adhesive tapes. We found that adhesion depended directly on shear force, and was independent of detachment angle. Therefore we reject the hypothesis that gecko toes peel like tape. The linear relation between adhesion and shear force is consistent with a critical angle of release in live gecko toes and isolated setal arrays, and also with our prior observations of single setae. We introduced a new model, frictional adhesion, for gecko pad attachment and compared it to existing models of adhesive contacts. In an analysis of clinging stability of a gecko on an inclined plane each adhesive model predicted a different force control strategy. The frictional adhesion model provides an explanation for the very low detachment forces observed in climbing geckos that does not depend on toe peeling.

The V-type H+ ATPase: molecular structure and function, physiological roles and regulation

Abstract: It was nearly 30 years before the V-type H+ ATPase was admitted to the small circle of bona fide transport ATPases alongside F-type and P-type ATPases. The V-type H+ ATPase is an ATP-driven enzyme that transforms the energy of ATP hydrolysis to electrochemical potential differences of protons across diverse biological membranes via the primary active transport of H+. In turn, the transmembrane electrochemical potential of H+ is used to drive a variety of (i) secondary active transport systems via H+-dependent symporters and antiporters and (ii) channel-mediated transport systems. For example, expression of Cl- channels or transporters next to the V-type H+ ATPase in vacuoles of plants and fungi and in lysosomes of animals brings about the acidification of the endosomal compartment, and the expression of the H+/neurotransmitter antiporter next to the V-type H+ ATPase concentrates neurotransmitters in synaptic vesicles. First found in association with endosomal membranes, the V-type H+ ATPase is now also found in increasing examples of plasma membranes where the proton pump energizes transport across cell membranes and entire epithelia. The molecular details reveal up to 14 protein subunits arranged in (i) a cytoplasmic V1 complex, which mediates the hydrolysis of ATP, and (ii) a membrane-embedded V0 complex, which translocates H+ across the membrane. Clever experiments have revealed the V-type H+ ATPase as a molecular motor akin to F-type ATPases. The hydrolysis of ATP turns a rotor consisting largely of one copy of subunits D and F of the V1 complex and a ring of six or more copies of subunit c of the V0 complex. The rotation of the ring is thought to deliver H+ from the cytoplasmic to the endosomal or extracellular side of the membrane, probably via channels formed by subunit a. The reversible dissociation of V1 and V0 complexes is one mechanism of physiological regulation that appears to be widely conserved from yeast to animal cells. Other mechanisms, such as subunit-subunit interactions or interactions of the V-type H+ ATPase with other proteins that serve physiological regulation, remain to be explored. Some diseases can now be attributed to genetic alterations of specific subunits of the V-type H+ ATPase.

Neuron–glia metabolic coupling and plasticity

Abstract: The coupling between synaptic activity and glucose utilization (neurometabolic coupling) is a central physiological principle of brain function that has provided the basis for 2-deoxyglucose-based functional imaging with positron emission tomography (PET). Astrocytes play a central role in neurometabolic coupling, and the basic mechanism involves glutamate-stimulated aerobic glycolysis; the sodium-coupled reuptake of glutamate by astrocytes and the ensuing activation of the Na-K-ATPase triggers glucose uptake and processing via glycolysis, resulting in the release of lactate from astrocytes. Lactate can then contribute to the activity-dependent fuelling of the neuronal energy demands associated with synaptic transmission. An operational model, the 'astrocyte-neuron lactate shuttle', is supported experimentally by a large body of evidence, which provides a molecular and cellular basis for interpreting data obtained from functional brain imaging studies. In addition, this neuron-glia metabolic coupling undergoes plastic adaptations in parallel with adaptive mechanisms that characterize synaptic plasticity. Thus, distinct subregions of the hippocampus are metabolically active at different time points during spatial learning tasks, suggesting that a type of metabolic plasticity, involving by definition neuron-glia coupling, occurs during learning. In addition, marked variations in the expression of genes involved in glial glycogen metabolism are observed during the sleep-wake cycle, with in particular a marked induction of expression of the gene encoding for protein targeting to glycogen (PTG) following sleep deprivation. These data suggest that glial metabolic plasticity is likely to be concomitant with synaptic plasticity.

Intraspecific variation in thermal tolerance and heat shock protein gene expression in common killifish, Fundulus heteroclitus.

Abstract: Populations of common killifish, Fundulus heteroclitus, are distributed along the Atlantic coast of North America through a steep latitudinal thermal gradient. We examined intraspecific variation in whole-animal thermal tolerance and its relationship to the heat shock response in killifish from the northern and southern extremes of the species range. Critical thermal maxima were significantly higher in southern than in northern fish by approximately 1.5 degrees C at a wide range of acclimation temperatures (from 2-34 degrees C), and critical thermal minima differed by approximately 1.5 degrees C at acclimation temperatures above 22 degrees C, converging on the freezing point of brackish water at lower acclimation temperatures. To determine whether these differences in whole-organism thermal tolerance were reflected in differences in either the sequence or regulation of the heat shock protein genes (hsps) we obtained complete cDNA sequences for hsc70, hsp70-1 and hsp70-2, and partial sequences of hsp90alpha and hsp90beta. There were no fixed differences in amino acid sequence between populations in either hsp70-1 or hsp70-2, and only a single conservative substitution between populations in hsc70. By contrast, there were significant differences between populations in the expression of many, but not all, of these genes. Both northern and southern killifish significantly increased hsp70-2 levels above control values (T(on)) at a heat shock temperature of 33 degrees C, but the magnitude of this induction was greater in northern fish, suggesting that northern fish may be more susceptible to thermal damage than are southern fish. In contrast, hsp70-1 mRNA levels increased gradually and to the same extent in response to heat shock in both populations. Hsc70 mRNA levels were significantly elevated by heat shock in southern fish, but not in northern fish. Similarly, the more thermotolerant southern killifish had a T(on) for hsp90alpha of 30 degrees C, 2 degrees C lower than that of northern fish. This observation combined with the ability of southern killifish to upregulate hsc70 in response to heat shock suggests a possible role for these hsps in whole-organism differences in thermal tolerance. These data highlight the importance of considering the complexity of the heat shock response across multiple isoforms when attempting to make linkages to whole-organism traits such as thermal tolerance.

Extreme diving of beaked whales

Abstract: Sound-and-orientation recording tags (DTAGs) were used to study 10 beaked whales of two poorly known species, Ziphius cavirostris (Zc) and Mesoplodon densirostris (Md). Acoustic behaviour in the deep foraging dives performed by both species (Zc: 28 dives by seven individuals; Md: 16 dives by three individuals) shows that they hunt by echolocation in deep water between 222 and 1885 m, attempting to capture about 30 prey/dive. This food source is so deep that the average foraging dives were deeper (Zc: 1070 m; Md: 835 m) and longer (Zc: 58 min; Md: 47 min) than reported for any other air-breathing species. A series of shallower dives, containing no indications of foraging, followed most deep foraging dives. The average interval between deep foraging dives was 63 min for Zc and 92 min for Md. This long an interval may be required for beaked whales to recover from an oxygen debt accrued in the deep foraging dives, which last about twice the estimated aerobic dive limit. Recent reports of gas emboli in beaked whales stranded during naval sonar exercises have led to the hypothesis that their deep-diving may make them especially vulnerable to decompression. Using current models of breath-hold diving, we infer that their natural diving behaviour is inconsistent with known problems of acute nitrogen supersaturation and embolism. If the assumptions of these models are correct for beaked whales, then possible decompression problems are more likely to result from an abnormal behavioural response to sonar.

Dietary choice behavior in Caenorhabditis elegans

Abstract: Animals have evolved diverse behaviors that serve the purpose of finding food in the environment. We investigated the food seeking strategy of the soil bacteria-eating nematode Caenorhabditis elegans. C. elegans bacterial food varies in quality: some species are easy to eat and support worm growth well, while others do not. We show that worms exhibit dietary choice: they hunt for high quality food and leave hard-to-eat bacteria. This food seeking behavior is enhanced in animals that have already experienced good food. When hunting for good food, worms alternate between two modes of locomotion, known as dwelling: movement with frequent stops and reversals; and roaming: straight rapid movement. On good food, roaming is very rare, while on bad food it is common. Using laser ablations and mutant analysis, we show that the AIY neurons serve to extend roaming periods, and are essential for efficient food seeking.

Dynamics of geckos running vertically

Abstract: Geckos with adhesive toe pads rapidly climb even smooth vertical surfaces. We challenged geckos (Hemidactylus garnotii) to climb up a smooth vertical track that contained a force platform. Geckos climbed vertically at up to 77·cm·s ‐1 with a stride frequency of 15·Hz using a trotting gait. During each step, whole body fore‐aft, lateral and normal forces all decreased to zero when the animal attached or detached its toe pads. Peak fore‐aft force was twice body weight at mid-step. Geckos climbed at a constant average velocity without generating decelerating forces on their center of mass in the direction of motion. Although mass-specific mechanical power to climb was ten times the value expected for level running, the total mechanical energy of climbing was only 5‐11% greater than the potential energy change. Fore- and hindlegs both pulled toward the midline, possibly loading the attachment mechanisms. Attachment and detachment of feet occupied 13% and 37% of stance time, respectively. As climbing speed increased, the absolute time required to attach and detach did not decrease, suggesting that the period of fore‐aft force production might be constrained. During ascent, the forelegs pulled toward, while hindlegs pushed away from the vertical surface, generating a net pitching moment toward the surface to counterbalance pitch-back away from the surface. Differential leg function appears essential for effective vertical as well as horizontal locomotion. Summary

Coordination of metabolic plasticity in skeletal muscle.

Abstract: Skeletal muscle is a highly malleable tissue, capable of pronounced metabolic and morphological adaptations in response to contractile activity (i.e. exercise). Each bout of contractile activity results in a coordinated alteration in the expression of a variety of nuclear DNA and mitochondrial DNA (mtDNA) gene products, leading to phenotypic adaptations. This results in an increase in muscle mitochondrial volume and changes in organelle composition, referred to as mitochondrial biogenesis. The functional consequence of this biogenesis is an improved resistance to fatigue. Signals initiated by the exercise bout involve changes in intracellular Ca2+ as well as alterations in energy status (i.e. ATP/ADP ratio) and the consequent activation of downstream kinases such as AMP kinase and Ca2+-calmodulin-activated kinases. These kinases activate transcription factors that bind DNA to affect the transcription of genes, the most evident manifestation of which occurs during the post-exercise recovery period when energy metabolism is directed toward anabolism, rather than contractile activity. An important protein that is affected by exercise is the transcriptional coactivator PGC-1alpha, which cooperates with multiple transcription factors to induce the expression of nuclear genes encoding mitochondrial proteins. Once translated in the cytosol, these mitochondrially destined proteins are imported into the mitochondrial outer membrane, inner membrane or matrix space via specific import machinery transport components. Contractile activity affects the expression of the import machinery, as well as the kinetics of import, thus facilitating the entry of newly synthesized proteins into the expanding organelle. An important set of proteins that are imported are the mtDNA transcription factors, which influence the expression and replication of mtDNA. While mtDNA contributes only 13 proteins to the synthesis of the organelle, these proteins are vital for the proper assembly of multi-subunit complexes of the respiratory chain, when combined with nuclear-encoded protein subunits. The expansion of skeletal muscle mitochondria during organelle biogenesis involves the assembly of an interconnected network system (i.e. a mitochondrial reticulum). This expansion of membrane size is influenced by the balance between mitochondrial fusion and fission. Thus, mitochondrial biogenesis is an adaptive process that requires the coordination of multiple cellular events, including the transcription of two genomes, the synthesis of lipids and proteins and the stoichiometric assembly of multisubunit protein complexes into a functional respiratory chain. Impairments at any step can lead to defective electron transport, a subsequent failure of ATP production and an inability to maintain energy homeostasis.

Simulations of optimized anguilliform swimming.

Abstract: The hydrodynamics of anguilliform swimming motions was investigated using three-dimensional simulations of the fluid flow past a self-propelled body. The motion of the body is not specified a priori, but is instead obtained through an evolutionary algorithm used to optimize the swimming efficiency and the burst swimming speed. The results of the present simulations support the hypothesis that anguilliform swimmers modify their kinematics according to different objectives and provide a quantitative analysis of the swimming motion and the forces experienced by the body. The kinematics of burst swimming is characterized by the large amplitude of the tail undulations while the anterior part of the body remains straight. In contrast, during efficient swimming behavior significant lateral undulation occurs along the entire length of the body. In turn, during burst swimming, the majority of the thrust is generated at the tail, whereas in the efficient swimming mode, in addition to the tail, the middle of the body contributes significantly to the thrust. The burst swimming velocity is 42% higher and the propulsive efficiency is 15% lower than the respective values during efficient swimming. The wake, for both swimming modes, consists largely of a double row of vortex rings with an axis aligned with the swimming direction. The vortex rings are responsible for producing lateral jets of fluid, which has been documented in prior experimental studies. We note that the primary wake vortices are qualitatively similar in both swimming modes except that the wake vortex rings are stronger and relatively more elongated in the fast swimming mode. The present results provide quantitative information of three-dimensional fluid-body interactions that may complement related experimental studies. In addition they enable a detailed quantitative analysis, which may be difficult to obtain experimentally, of the different swimming modes linking the kinematics of the motion with the forces acting on the self-propelled body. Finally, the optimization procedure helps to identify, in a systematic fashion, links between swimming motion and biological function.

Environment and plasticity of myogenesis in teleost fish.

Abstract: Embryonic development in teleosts is profoundly affected by environmental conditions, particularly temperature and dissolved oxygen concentrations. The environment determines the rate of myogenesis, the composition of sub-cellular organelles, patterns of gene expression, and the number and size distribution of muscle fibres. During the embryonic and larval stages, muscle plasticity to the environment is usually irreversible due to the rapid pace of ontogenetic change. In the early life stages, muscle can affect locomotory performance and behaviour, with potential consequences for larval survival. Postembryonic growth involves myogenic progenitor cells (MPCs) that originate in the embryo. The embryonic temperature regime can have long-term consequences for the growth of skeletal muscle in some species, including the duration and intensity of myotube formation in adult stages. In juvenile and adult fish, abiotic (temperature, day-length, water flow characteristics, hypoxia) and biotic factors (food availability, parasitic infection) have complex effects on the signalling pathways regulating the proliferation and differentiation of MPCs, protein synthesis and degradation, and patterns of gene expression. The phenotypic responses observed to the environment frequently vary during ontogeny and are integrated with endogenous physiological rhythms, particularly sexual maturation. Studies with model teleosts provide opportunities for investigating the underlying genetic mechanisms of muscle plasticity that can subsequently be applied to non-model species of more ecological or commercial interest.

Interactions between the human gastrocnemius muscle and the Achilles tendon during incline, level and decline locomotion

Abstract: Muscles are required to perform or absorb mechanical work under different conditions. However the ability of a muscle to do this depends on the interaction between its contractile components and its elastic components. In the present study we have used ultrasound to examine the length changes of the gastrocnemius medialis muscle fascicle along with those of the elastic Achilles tendon during locomotion under different incline conditions. Six male participants walked (at 5 km h-1) on a treadmill at grades of -10%, 0% and 10% and ran (at 10 km h-1) at grades of 0% and 10%, whilst simultaneous ultrasound, electromyography and kinematics were recorded. In both walking and running, force was developed isometrically; however, increases in incline increased the muscle fascicle length at which force was developed. Force was developed at shorter muscle lengths for running when compared to walking. Substantial levels of Achilles tendon strain were recorded in both walking and running conditions, which allowed the muscle fascicles to act at speeds more favourable for power production. In all conditions, positive work was performed by the muscle. The measurements suggest that there is very little change in the function of the muscle fascicles at different slopes or speeds, despite changes in the required external work. This may be a consequence of the role of this biarticular muscle or of the load sharing between the other muscles of the triceps surae.

Phenotypic plasticity and experimental evolution

Abstract: Natural or artificial selection that favors higher values of a particular trait within a given population should engender an evolutionary response that increases the mean value of the trait. For this prediction to hold, the phenotypic variance of the trait must be caused in part by additive effects of alleles segregating in the population, and also the trait must not be too strongly genetically correlated with other traits that are under selection. Another prediction, rarely discussed in the literature, is that directional selection should favor alleles that increase phenotypic plasticity in the direction of selection, where phenotypic plasticity is defined as the ability of one genotype to produce more than one phenotype when exposed to different environments. This prediction has received relatively little empirical attention. Nonetheless, many laboratory experiments impose selection regimes that could allow for the evolution of enhanced plasticity (e.g. desiccation trials with Drosophila that last for several hours or days). We review one example that involved culturing of Drosophila on lemon for multiple generations and then tested for enhanced plasticity of detoxifying enzymes. We also review an example with vertebrates that involves selective breeding for high voluntary activity levels in house mice, targeting wheel-running behavior on days 5+6 of a 6-day wheel exposure. This selection regime allows for the possibility of wheel running itself or subordinate traits that support such running to increase in plasticity over days 1-4 of wheel access. Indeed, some traits, such as the concentration of the glucose transporter GLUT4 in gastrocnemius muscle, do show enhanced plasticity in the selected lines over a 5-6 day period. In several experiments we have housed mice from both the Selected (S) and Control (C) lines with or without wheel access for several weeks to test for differences in plasticity (training effects). A variety of patterns were observed, including no training effects in either S or C mice, similar changes in both the S and C lines, greater changes in the S lines but in the same direction in the C lines, and even opposite directions of change in the S and C lines. For some of the traits that show a greater training effect in the S lines, but in the same direction as in C lines, the greater effect can be explained statistically by the greater wheel running exhibited by S lines ('more pain, more gain'). For others, however, the differences seem to reflect inherently greater plasticity in the S lines (i.e. for a given amount of stimulus, such as wheel running/day, individuals in the S lines show a greater response as compared with individuals in the C lines). We suggest that any selection experiment in which the selective event is more than instantaneous should explore whether plasticity in the appropriate (adaptive) direction has increased as a component of the response to selection.

Constraints of tolerance: why are desiccation-tolerant organisms so small or rare?

Abstract: Drying to equilibrium with the air kills nearly all animals and flowering plants, including livestock and crops. This makes drought a key ecological problem for terrestrial life and a major cause of human famine. However, the ability to tolerate complete desiccation is widespread in organisms that are either <5 mm long or found mainly where desiccation-sensitive organisms are scarce. This suggests that there is a trade-off between desiccation tolerance and growth. Recent molecular and biochemical research shows that organisms tolerate desiccation through a set of mechanisms, including sugars that replace water and form glasses, proteins that stabilize macromolecules and membranes, and anti-oxidants that counter damage by reactive oxygen species. These protections are often induced by drying, and some of the genes involved may be homologous in microbes, plants and animals. Understanding how mechanisms of desiccation tolerance may constrain growth might show how to undo the constraint in some economically important macroorganisms and elucidate the much-studied but elusive relationship between tolerance of stress and productivity.

Effective elastic modulus of isolated gecko setal arrays

Abstract: Conventional pressure sensitive adhesives (PSAs) are fabricated from soft viscoelastic materials that satisfy Dahlquist's criterion for tack with a Young's modulus (E) of 100 kPa or less at room temperature and 1 Hz. In contrast, the adhesive on the toes of geckos is made of beta-keratin, a stiff material with E at least four orders of magnitude greater than the upper limit of Dahlquist's criterion. Therefore, one would not expect a beta-keratin structure to function as a PSA by deforming readily to make intimate molecular contact with a variety of surface profiles. However, since the gecko adhesive is a microstructure in the form of an array of millions of high aspect ratio shafts (setae), the effective elastic modulus (E(eff)) is much lower than E of bulk beta-keratin. In the first test of the E(eff) of a gecko setal adhesive, we measured the forces resulting from deformation of isolated arrays of tokay gecko (Gekko gecko) setae during vertical compression, and during tangential compression at angles of +45 degrees and -45 degrees . We tested the hypothesis that E(eff) of gecko setae falls within Dahlquist's criterion for tack, and evaluated the validity of a model of setae as cantilever beams. Highly linear forces of deformation under all compression conditions support the cantilever model. E(eff) of setal arrays during vertical and +45 degrees compression (along the natural path of drag of the setae) were 83+/-4.0 kPa and 86+/-4.4 kPa (means +/- s.e.m.), respectively. Consistent with the predictions of the cantilever model, setae became significantly stiffer when compressed against the natural path of drag: E(eff) during -45 degrees compression was 110+/-4.7 kPa. Unlike synthetic PSAs, setal arrays act as Hookean elastic solids; setal arrays function as a bed of springs with a directional stiffness, assisting alignment of the adhesive spatular tips with the contact surface during shear loading.

Effect of temperature on maximum swimming speed and cost of transport in juvenile European sea bass (Dicentrarchus labrax).

Abstract: This study is an attempt to gain an integrated understanding of the interactions between temperature, locomotion activity and metabolism in the European sea bass ( Dicentrarchus labrax ). To our knowledge this study is among the few that have investigated the influence of the seasonal changes in water temperature on swimming performance in fish. Using a Brett-type swim-tunnel respirometer the relationship between oxygen consumption and swimming speed was determined in fish acclimatised to 7, 11, 14, 18, 22, 26 and 30°C. The corresponding maximum swimming speed ( U max), optimal swimming speed ( U opt), active (AMR) and standard (SMR) metabolic rates as well as aerobic metabolic scope (MS) were calculated. Using simple mathematical functions, these parameters were modelled as a function of water temperature and swimming speed. Both SMR and AMR were positively related to water temperature up to 24°C. Above 24°C SMR and AMR levelled off and MS tended to decrease. We found a tight relationship between AMR and U max and observed that raising the temperature increased AMR and increased swimming ability. However, although fish swam faster at high temperature, the net cost of transport (COTnet) at a given speed was not influence by the elevation of the water temperature. Although U opt doubled between 7°C and 30°C (from 0.3 to 0.6 m s-1), metabolic rate at U opt represented a relatively constant fraction of the animal active metabolic rate (40-45%). A proposed model integrates the effects of water temperature on the interaction between metabolism and swimming performance. In particular the controlling effect of temperature on AMR is shown to be the key factor limiting maximal swimming speed of sea bass.

Functional, structural and molecular plasticity of mammalian skeletal muscle in response to exercise stimuli

Abstract: Biological systems have acquired effective adaptive strategies to cope with physiological challenges and to maximize biochemical processes under imposed constraints. Striated muscle tissue demonstrates a remarkable malleability and can adjust its metabolic and contractile makeup in response to alterations in functional demands. Activity-dependent muscle plasticity therefore represents a unique model to investigate the regulatory machinery underlying phenotypic adaptations in a fully differentiated tissue. Adjustments in form and function of mammalian muscle have so far been characterized at a descriptive level, and several major themes have evolved. These imply that mechanical, metabolic and neuronal perturbations in recruited muscle groups relay to the specific processes being activated by the complex physiological stimulus of exercise. The important relationship between the phenotypic stimuli and consequent muscular modifications is reflected by coordinated differences at the transcript level that match structural and functional adjustments in the new training steady state. Permanent alterations of gene expression thus represent a major strategy for the integration of phenotypic stimuli into remodeling of muscle makeup. A unifying theory on the molecular mechanism that connects the single exercise stimulus to the multi-faceted adjustments made after the repeated impact of the muscular stress remains elusive. Recently, master switches have been recognized that sense and transduce the individual physical and chemical perturbations induced by physiological challenges via signaling cascades to downstream gene expression events. Molecular observations on signaling systems also extend the long-known evidence for desensitization of the muscle response to endurance exercise after the repeated impact of the stimulus that occurs with training. Integrative approaches involving the manipulation of single factors and the systematic monitoring of downstream effects at multiple levels would appear to be the ultimate method for pinpointing the mechanism of muscle remodeling. The identification of the basic relationships underlying the malleability of muscle tissue is likely to be of relevance for our understanding of compensatory processes in other tissues, species and organisms.

Following the heart: temperature and salinity effects on heart rate in native and invasive species of blue mussels (genus Mytilus).

Abstract: The three species of blue mussels, Mytilus trossulus Gould 1850, M. edulis Linnaeus 1758 and M. galloprovincialis Lamarck 1819, have distinct global distribution patterns that are hypothesized to reflect differences in their tolerances of temperature and salinity. We examined effects on heart rate (beats min–1) of acute exposure and acclimation to different combinations of temperature and salinity to test this hypothesis and, in the context of the invasive success of M. galloprovincialis , to gain insights into the factors that may explain the replacement of the temperate Pacific native, M. trossulus , by this Mediterranean Sea invader along much of the California coast. Heart rate of M. trossulus was significantly higher than that of M. galloprovincialis , consistent with evolutionary adaptation to a lower habitat temperature (temperature compensation) in the former species. Heart rates of M. trossulus / M. galloprovincialis hybrids were intermediate between those of the parental species. Following acclimation to 14°C and 21°C, heart rates of all species exhibited partial compensation to temperature. Heart rate increased with rising temperature until a high temperature was reached at which point activity fell sharply, the high critical temperature ( H crit). H crit increased with increasing acclimation temperature and differed among species in a pattern that reflected their probable evolutionary adaptation temperatures: M. galloprovincialis is more heat tolerant than the other two congeners. Ability to sustain heart function in the cold also reflected evolutionary history: M. trossulus is more cold tolerant than M. galloprovincialis . Heart rates for all three congeners decreased gradually in response to acute reductions in salinity until a low salinity ( S crit) was reached at which heart rate dropped precipitously. S crit decreased with decreasing salinity of acclimation and was generally lowest for M. galloprovincialis . Mortality during acclimation under common garden conditions was greatest in M. trossulus and was highest at high acclimation temperatures and salinities. These intrinsic differences in basal heart rate, thermal and salinity responses, acclimatory capacity, and survivorship are discussed in the contexts of the species' biogeographic patterning and, for the invasive species M. galloprovincialis , the potential for further range expansion along the Pacific coast of North America.

The advantages of a rolling foot in human walking

Abstract: The plantigrade human foot rolls over the ground during each walking step, roughly analogous to a wheel. The center of pressure progresses on the ground like a wheel of radius 0.3 L (leg length). We examined the effect of varying foot curvature on the mechanics and energetics of walking. We controlled curvature by attaching rigid arc shapes of various radii to the bottoms of rigid boots restricting ankle motion. We measured mechanical work performed on the center of mass (COM), and net metabolic rate, in human subjects (N=10) walking with seven arc radii from 0.02-0.40 m. Simple models of dynamic walking predict that redirection of COM velocity requires step-to-step transition work, decreasing quadratically with arc radius. Metabolic cost would be expected to change in proportion to mechanical work. We measured the average rate of negative work performed on the COM, and found that it followed the trend well (r2=0.95), with 2.37 times as much work for small radii as for large. Net metabolic rate (subtracting quiet standing) also decreased with increasing arc radius to a minimum at 0.3 L, with a slight increase thereafter. Maximum net metabolic rate was 6.25 W kg(-1) (for small-radius arc feet), about 59% greater than the minimum rate of 3.93 W kg(-1), which in turn was about 45% greater than the rate in normal walking. Metabolic rate was fit reasonably well (r2=0.86) by a quadratic curve, but exceeded that expected from COM work for extreme arc sizes. Other factors appear to increase metabolic cost for walking on very small and very large arc feet. These factors may include effort expended to stabilize the joints (especially the knee) or to maintain balance. Rolling feet with curvature 0.3 L appear energetically advantageous for plantigrade walking, partially due to decreased work for step-to-step transitions.

Hydrogen sulfide as an oxygen sensor/transducer in vertebrate hypoxic vasoconstriction and hypoxic vasodilation.

Abstract: How vertebrate blood vessels sense acute hypoxia and respond either by constricting (hypoxic vasoconstriction) or dilating (hypoxic vasodilation) has not been resolved. In the present study we compared the mechanical and electrical responses of select blood vessels to hypoxia and H2S, measured vascular H2S production, and evaluated the effects of inhibitors of H2S synthesis and addition of the H2S precursor, cysteine, on hypoxic vasoconstriction and hypoxic vasodilation. We found that: (1) in all vertebrate vessels examined to date, hypoxia and H2S produce temporally and quantitatively identical responses even though the responses vary from constriction (lamprey dorsal aorta; lDA), to dilation (rat aorta; rA), to multi-phasic (rat and bovine pulmonary arteries; rPA and bPA, respectively). (2) The responses of lDA, rA and bPA to hypoxia and H2S appear competitive; in the presence of one stimulus, the response to the other stimulus is substantially or completely eliminated. (3) Hypoxia and H2S produce the same degree of cell depolarization in bPA. (4) H2S is constitutively synthesized by lDA and bPA vascular smooth muscle. (5) Inhibition of H2S synthesis inhibits the hypoxic response of lDA, rA, rPA and bPA. (6) Addition of the H2S precursor, cysteine, doubles hypoxic contraction in lDA, prolongs contraction in bPA and alters the re-oxygenation response of rA. These studies suggest that H2S may serve as an O2 sensor/transducer in the vascular responses to hypoxia. In this model, the concentration of vasoactive H2S in the vessel is governed by the balance between endogenous H2S production and its oxidation by available O2.

Kinematics of foraging dives and lunge-feeding in fin whales

Abstract: Fin whales are among the largest predators on earth, yet little is known about their foraging behavior at depth. These whales obtain their prey by lunge-feeding, an extraordinary biomechanical event where large amounts of water and prey are engulfed and filtered. This process entails a high energetic cost that effectively decreases dive duration and increases post-dive recovery time. To examine the body mechanics of fin whales during foraging dives we attached high-resolution digital tags, equipped with a hydrophone, a depth gauge and a dual-axis accelerometer, to the backs of surfacing fin whales in the Southern California Bight. Body pitch and roll were estimated by changes in static gravitational acceleration detected by orthogonal axes of the accelerometer, while higher frequency, smaller amplitude oscillations in the accelerometer signals were interpreted as bouts of active fluking. Instantaneous velocity of the whale was determined from the magnitude of turbulent flow noise measured by the hydrophone and confirmed by kinematic analysis. Fin whales employed gliding gaits during descent, executed a series of lunges at depth and ascended to the surface by steady fluking. Our examination of body kinematics at depth reveals variable lunge-feeding behavior in the context of distinct kinematic modes, which exhibit temporal coordination of rotational torques with translational accelerations. Maximum swimming speeds during lunges match previous estimates of the flow-induced pressure needed to completely expand the buccal cavity during feeding.

The evolutionary consequences of ecological interactions mediated through phenotypic plasticity

Abstract: Phenotypic plasticity describes the capacity of a genotype to exhibit a range of phenotypes in response to variation in the environment. Environmental variation encompasses both abiotic and biotic components of the environment, including interactions among organisms. The strength and outcome of many ecological interactions, ranging from antagonism to mutualism, are mediated through the phenotypically plastic responses of one or more players in the interaction. Herein, three broadly defined, non-mutually exclusive, evolutionary consequences of ecological interactions mediated through phenotypic plasticity are discussed. (1) The predictable plastic response of one partner can favor behaviors, physiological responses, and life history traits of an interacting partner that manipulate, circumvent, or ameliorate the response of that partner. (2) Phenotypic plasticity can generate substantial spatial and temporal variation within and among populations. Such phenotypic variation can depend on the density and identity of interacting players in an ecological community, and can ultimately affect the evolutionary outcome of ecological interactions. (3) Phenotypic plasticity affects the strength and direction of natural selection. Ecological interactions mediated through phenotypic plasticity are ubiquitous in nature, and the potential evolutionary consequences of these interactions illustrate the complexity inherent in understanding evolution in a community context.

When bad things happen to good fish: the loss of hemoglobin and myoglobin expression in Antarctic icefishes

Abstract: The Antarctic icefishes (Family Channichthyidae) provide excellent examples of unique traits that can arise in a chronically cold and isolated environment. Their loss of hemoglobin (Hb) expression, and in some cases, loss of myoglobin (Mb) expression, has taught us much about the function of these proteins. Although absences of the proteins are fixed traits in icefishes, the losses do not appear to be of adaptive value. Contrary to some suggestions, loss of Hb has led to higher energetic costs for circulating blood, and losses of Mb have reduced cardiac performance. Moreover, losses of Hb and Mb have resulted in extensive modifications to the cardiovascular system to ensure adequate oxygen delivery to working muscles. Recent studies suggest that losses of Hb and Mb, and their associated nitric oxide (NO)-oxygenase activities, may have accelerated the development and evolution of these cardiovascular modifications. The high levels of NO that should occur in the absence of Hb and Mb have been shown in other animal groups to lead to an increase in tissue vascularization, an increase in the lumenal diameter of blood vessels, and an increase in mitochondrial densities. These characteristics are all hallmark traits of Antarctic icefishes. Homeostatic feedback mechanisms thus may have accelerated evolution of the pronounced cardiovascular traits of Antarctic icefishes.

Intestinal anion exchange in marine fish osmoregulation

Abstract: Despite early reports, dating back three quarters of a century, of high total CO(2) concentrations in the intestinal fluids of marine teleost fishes, only the past decade has provided some insight into the functional significance of this phenomenon. It is now being recognized that intestinal anion exchange is responsible for high luminal HCO(3)(-) and CO(3)(2-) concentrations while at the same time contributing substantially to intestinal Cl(-) and thereby water absorption, which is vital for marine fish osmoregulation. In species examined to date, the majority of HCO(3)(-) secreted by the apical anion exchange process is derived from hydration of metabolic CO(2) with the resulting H(+) being extruded via a Na(+):H(+) exchange mechanism in the basolateral membrane. The basolateral H(+) extrusion is critical for the apical anion exchange and relies on the Na(+) gradient established by the Na(+)-K(+)-ATPase. This enzyme thereby ultimately fuels the secondary active transport of HCO(3)(-) and Cl(-) by the apical anion exchanger. High cellular HCO(3)(-) concentrations (>10 mmol l(-1)) are required for the anion exchange process and could be the result of both a high metabolic activity of the intestinal epithelium and a close association of the anion exchange protein and the enzyme carbonic anhydrase. The anion exchange activity in vivo is likely most pronounced in the anterior segment and results in net intestinal acid absorption. In contrast to other water absorbing vertebrate epithelia, the marine teleost intestine absorbs what appears to be a hypertonic fluid to displace diffusive fluid loss to the marine environment.

Unifying constructal theory for scale effects in running, swimming and flying

Abstract: SUMMARY Biologists have treated the view that fundamental differences exist between running, flying and swimming as evident, because the forms of locomotion and the animals are so different: limbs and wings vs body undulations, neutrally buoyant vs weighted bodies, etc. Here we show that all forms of locomotion can be described by a single physics theory. The theory is an invocation of the principle that flow systems evolve in such a way that they destroy minimum useful energy (exergy, food). This optimization approach delivers in surprisingly direct fashion the observed relations between speed and body mass ( M b ) raised to 1/6, and between frequency (stride, flapping) and \(M_{\mathrm{b}}^{-1{/}6}\) , and shows why these relations hold for running, flying and swimming. Animal locomotion is an optimized two-step intermittency: an optimal balance is achieved between the vertical loss of useful energy (lifting the body weight, which later drops), and the horizontal loss caused by friction against the surrounding medium. The theory predicts additional features of animal design: the Strouhal number constant, which holds for running as well as flying and swimming, the proportionality between force output and mass in animal motors, and the fact that undulating swimming and flapping flight occur only if the body Reynolds number exceeds approximately 30. This theory, and the general body of work known as constructal theory, together now show that animal movement (running, flying, swimming) and fluid eddy movement (turbulent structure) are both forms of optimized intermittent movement.

Silken toolkits: biomechanics of silk fibers spun by the orb web spider Argiope argentata (Fabricius 1775).

Abstract: Orb-weaving spiders spin five fibrous silks from differentiated glands that contain unique sets of proteins Despite diverse ecological functions, the mechanical properties of most of these silks are not well characterized Here, we quantify the mechanical performance of this toolkit of silks for the silver garden spider Argiope argentata Four silks exhibit viscoelastic behaviour typical of polymers, but differ statistically from each other by up to 250% in performance, giving each silk a distinctive suite of material properties Major ampullate silk is 50% stronger than other fibers, but also less extensible Aciniform silk is almost twice as tough as other silks because of high strength and extensibility Capture spiral silk, coated with aqueous glue, is an order of magnitude stretchier than other silks Dynamic mechanical properties are qualitatively similar, but quantitatively vary by up to 300% among silks Storage moduli are initially nearly constant and increase after fiber yield, whereas loss tangents reach maxima of 01-02 at the yield The remarkable mechanical diversity of Argiope argentata silks probably results in part from the different molecular structures of fibers and can be related to the specific ecological role of each silk Our study indicates substantial potential to customize the mechanics of bioengineered silks

The cellular response to heat stress in the goby Gillichthys mirabilis: a cDNA microarray and protein-level analysis.

Abstract: The cellular response to stress relies on the rapid induction of genes encoding proteins involved in preventing and repairing macromolecular damage incurred as a consequence of environmental insult. To increase our understanding of the scope of this response, a cDNA microarray, consisting of 9207 cDNA clones, was used to monitor gene expression changes in the gill and white muscle tissues of a eurythermic fish, Gillichthys mirabilis (Gobiidae) exposed to ecologically relevant heat stress. In each tissue, the induction or repression of over 200 genes was observed. These genes are associated with numerous biological processes, including the maintenance of protein homeostasis, cell cycle control, cytoskeletal reorganization, metabolic regulation and signal transduction, among many others. In both tissues, the molecular chaperones, certain transcription factors and a set of additional genes with various functions were induced in a similar manner; however, the majority of genes displayed tissue-specific responses. In gill, thermal stress induced the expression of the major structural components of the cytoskeleton, whereas these same genes did not respond to heat in muscle. In muscle, many genes involved in promoting cell growth and proliferation were repressed, perhaps to conserve energy for repair and replacement of damaged macromolecules, but a similar repression was not observed in the gill. Many of the observed changes in gene expression were similar to those described in model species whereas many others were unexpected. Measurements of the concentrations of the protein products of selected genes revealed that in each case an induction in mRNA synthesis correlated with an increase in protein production, though the timing and magnitude of the increase in protein was not consistently predicted by mRNA concentration, an important consideration in assessing the condition of the stressed cell using transcriptomic analysis.

Water relations of tetrapod integument

Abstract: The vertebrate integument represents an evolutionary compromise between the needs for mechanical protection and those of sensing the environment and regulating the exchange of materials and energy. Fibrous keratins evolved as a means of strengthening the integument while simultaneously providing a structural support for lipids, which comprise the principal barrier to cutaneous water efflux in terrestrial taxa. Whereas lipids are of fundamental importance to water barriers, the efficacy of these barriers depends in many cases on structural features that enhance or maintain the integrity of function. Amphibians are exceptional among tetrapods in having very little keratin and a thin stratum corneum. Thus, effective lipid barriers that are present in some specialized anurans living in xeric habitats are external to the epidermis, whereas lipid barriers of amniotes exist as a lipid-keratin complex within the stratum corneum. Amphibians prevent desiccation of the epidermis and underlying tissues either by evaporating water from a superficial aqueous film, which must be replenished, or by shielding the stratum corneum with superficial lipids. Water barrier function in vertebrates generally appears to be relatively fixed, although various species have ;plasticity' to adjust the barrier effectiveness facultatively. While it is clear that both phenotypic plasticity and genetic adaptation can account for covariation between environment and skin resistance to water efflux, studies of the relative importance of these two phenomena are few. Fundamental mechanisms for adjusting the skin water barrier include changes in barrier thickness, composition and physicochemical properties of cutaneous lipids, and/or geometry of the barrier within the epidermis. While cutaneous lipids have been studied extensively in the contexts of disease and cosmetics, relatively little is known about the processes of permeability barrier ontogenesis related to adaptation and environment. Advances in such knowledge have didactic significance for understanding vertebrate evolution as well as practical application to clinical dermatology.

Influence of the muscle-tendon unit's mechanical and morphological properties on running economy

Abstract: The purpose of this study was to test the hypothesis that runners having different running economies show differences in the mechanical and morphological properties of their muscle-tendon units (MTU) in the lower extremities. Twenty eight long-distance runners (body mass: 76.8+/-6.7 kg, height: 182+/-6 cm, age: 28.1+/-4.5 years) participated in the study. The subjects ran on a treadmill at three velocities (3.0, 3.5 and 4.0 m s(-1)) for 15 min each. The V(O(2)) consumption was measured by spirometry. At all three examined velocities the kinematics of the left leg were captured whilst running on the treadmill using a high-speed digital video camera operating at 250 Hz. Furthermore the runners performed isometric maximal voluntary plantarflexion and knee extension contractions at eleven different MTU lengths with their left leg on a dynamometer. The distal aponeuroses of the gastrocnemius medialis (GM) and vastus lateralis (VL) were visualised by ultrasound during plantarflexion and knee extension, respectively. The morphological properties of the GM and VL (fascicle length, angle of pennation, and thickness) were determined at three different lengths for each MTU. A cluster analysis was used to classify the subjects into three groups according to their V(O(2)) consumption at all three velocities (high running economy, N=10; moderate running economy, N=12; low running economy, N=6). Neither the kinematic parameters nor the morphological properties of the GM and VL showed significant differences between groups. The most economical runners showed a higher contractile strength and a higher normalised tendon stiffness (relationship between tendon force and tendon strain) in the triceps surae MTU and a higher compliance of the quadriceps tendon and aponeurosis at low level tendon forces. It is suggested that at low level forces the more compliant quadriceps tendon and aponeurosis will increase the force potential of the muscle while running and therefore the volume of active muscle at a given force generation will decrease.

Foraging Blainville's beaked whales (Mesoplodon densirostris) produce distinct click types matched to different phases of echolocation.

Abstract: SUMMARY Blainville9s beaked whales ( Mesoplodon densirostris Blainville) echolocate for prey during deep foraging dives. Here we use acoustic tags to demonstrate that these whales, in contrast to other toothed whales studied, produce two distinct types of click sounds during different phases in biosonar-based foraging. Search clicks are emitted during foraging dives with inter-click intervals typically between 0.2 and 0.4 s. They have the distinctive form of an FM upsweep (modulation rate of about 110 kHz ms -1 ) with a -10 dB bandwidth from 26 to 51 kHz and a pulse length of 270 μs, somewhat similar to chirp signals in bats and Cuvier9s beaked whales ( Ziphius cavirostris Cuvier), but quite different from clicks of other toothed whales studied. In comparison, the buzz clicks, produced in short bursts during the final stage of prey capture, are short (105 μs) transients with no FM structure and a -10 dB bandwidth from 25 to 80 kHz or higher. Buzz clicks have properties similar to clicks reported from large delphinids and hold the potential for higher temporal resolution than the FM clicks. It is suggested that the two click types are adapted to the separate problems of target detection and classification versus capture of low target strength prey in a cluttered acoustic environment.