Showing papers in "The Journal of Experimental Biology in 2010"

The physiology of climate change: how potentials for acclimatization and genetic adaptation will determine 'winners' and 'losers'.

Abstract: SUMMARY Physiological studies can help predict effects of climate change through determining which species currently live closest to their upper thermal tolerance limits, which physiological systems set these limits, and how species differ in acclimatization capacities for modifying their thermal tolerances. Reductionist studies at the molecular level can contribute to this analysis by revealing how much change in sequence is needed to adapt proteins to warmer temperatures — thus providing insights into potential rates of adaptive evolution — and determining how the contents of genomes — protein-coding genes and gene regulatory mechanisms — influence capacities for adapting to acute and long-term increases in temperature. Studies of congeneric invertebrates from thermally stressful rocky intertidal habitats have shown that warm-adapted congeners are most susceptible to local extinctions because their acute upper thermal limits (LT 50 values) lie near current thermal maxima and their abilities to increase thermal tolerance through acclimation are limited. Collapse of cardiac function may underlie acute and longer-term thermal limits. Local extinctions from heat death may be offset by in-migration of genetically warm-adapted conspecifics from mid-latitude ‘hot spots’, where midday low tides in summer select for heat tolerance. A single amino acid replacement is sufficient to adapt a protein to a new thermal range. More challenging to adaptive evolution are lesions in genomes of stenotherms like Antarctic marine ectotherms, which have lost protein-coding genes and gene regulatory mechanisms needed for coping with rising temperature. These extreme stenotherms, along with warm-adapted eurytherms living near their thermal limits, may be the major ‘losers’ from climate change.

Oxygen- and capacity-limitation of thermal tolerance: a matrix for integrating climate-related stressor effects in marine ecosystems.

Abstract: SUMMARY The concept of oxygen- and capacity-dependent thermal tolerance in aquatic ectotherms has successfully explained climate-induced effects of rising temperatures on species abundance in the field. Oxygen supply to tissues and the resulting aerobic performance characters thus form a primary link between organismal fitness and its role and functioning at the ecosystem level. The thermal window of performance in water breathers matches their window of aerobic scope. Loss of performance reflects the earliest level of thermal stress, caused by hypoxaemia and the progressive mismatch of oxygen supply and demand at the borders of the thermal envelope. Oxygen deficiency elicits the transition to passive tolerance and associated systemic and cellular stress signals like hormonal responses or oxidative stress as well as the use of protection mechanisms like heat shock proteins at thermal extremes. Thermal acclimatization between seasons or adaptation to a climate regime involves shifting thermal windows and adjusting window widths. The need to specialize on a limited temperature range results from temperature-dependent trade-offs at several hierarchical levels, from molecular structure to whole-organism functioning, and may also support maximized energy efficiency. Various environmental factors like CO 2 (ocean acidification) and hypoxia interact with these principal relationships. Existing knowledge suggests that these factors elicit metabolic depression supporting passive tolerance to thermal extremes. However, they also exacerbate hypoxaemia, causing a narrowing of thermal performance windows and prematurely leading the organism to the limits of its thermal acclimation capacity. The conceptual analysis suggests that the relationships between energy turnover, the capacities of activity and other functions and the width of thermal windows may lead to an integrative understanding of specialization on climate and, as a thermal matrix, of sensitivity to climate change and the factors involved. Such functional relationships might also relate to climate-induced changes in species interactions and, thus, community responses at the ecosystem level.

Insect overwintering in a changing climate

Abstract: Insects are highly successful animals inhabiting marine, freshwater and terrestrial habitats from the equator to the poles. As a group, insects have limited ability to regulate their body temperature and have thus required a range of strategies to support life in thermally stressful environments, including behavioural avoidance through migration and seasonal changes in cold tolerance. With respect to overwintering strategies, insects have traditionally been divided into two main groups: freeze tolerant and freeze avoiding, although this simple classification is underpinned by a complex of interacting processes, i.e. synthesis of ice nucleating agents, cryoprotectants, antifreeze proteins and changes in membrane lipid composition. Also, in temperate and colder climates, the overwintering ability of many species is closely linked to the diapause state, which often increases cold tolerance ahead of temperature-induced seasonal acclimatisation. Importantly, even though most species can invoke one or both of these responses, the majority of insects die from the effects of cold rather than freezing. Most studies on the effects of a changing climate on insects have focused on processes that occur predominantly in summer (development, reproduction) and on changes in distributions rather than winter survival per se. For species that routinely experience cold stress, a general hypothesis would be that predicted temperature increases of 1 degree C to 5 degrees C over the next 50-100 years would increase winter survival in some climatic zones. However, this is unlikely to be a universal effect. Negative impacts may occur if climate warming leads to a reduction or loss of winter snow cover in polar and sub-polar areas, resulting in exposure to more severe air temperatures, increasing frequency of freeze-thaw cycles and risks of ice encasement. Likewise, whilst the dominant diapause-inducing cue (photoperiod) will be unaffected by global climate change, higher temperatures may modify normal rates of development, leading to a decoupling of synchrony between diapause-sensitive life-cycle stages and critical photoperiods for diapause induction. In terms of climate warming and potential heat stress, the most recent predictions of summer temperatures in Europe of 40 degrees C or higher in 50-75 years, are close to the current upper lethal limit of some insects. Long-term data sets on insect distributions and the timing of annual migrations provide strong evidence for 'positive' responses to higher winter temperatures over timescales of the past 20-50 years in North America, Europe and Asia.

Variation in the heat shock response and its implication for predicting the effect of global climate change on species' biogeographical distribution ranges and metabolic costs

Abstract: The preferential synthesis of heat shock proteins (Hsps) in response to thermal stress [the heat shock response (HSR)] has been shown to vary in species that occupy different thermal environments. A survey of case studies of aquatic (mostly marine) organisms occupying stable thermal environments at all latitudes, from polar to tropical, shows that they do not in general respond to heat stress with an inducible HSR. Organisms that occupy highly variable thermal environments (variations up to >20 degrees C), like the intertidal zone, induce the HSR frequently and within the range of body temperatures they normally experience, suggesting that the response is part of their biochemical strategy to occupy this thermal niche. The highest temperatures at which these organisms can synthesize Hsps are only a few degrees Celsius higher than the highest body temperatures they experience. Thus, they live close to their thermal limits and any further increase in temperature is probably going to push them beyond those limits. In comparison, organisms occupying moderately variable thermal environments (<10 degrees C), like the subtidal zone, activate the HSR at temperatures above those they normally experience in their habitats. They have a wider temperature range above their body temperature range over which they can synthesize Hsps. Contrary to our expectations, species from highly (in comparison with moderately) variable thermal environments have a limited acclimatory plasticity. Due to this variation in the HSR, species from stable and highly variable environments are likely to be more affected by climate change than species from moderately variable environments.

The cause of global amphibian declines: a developmental endocrinologist's perspective

Abstract: Greater than 70% of the world's amphibian species are in decline. We propose that there is probably not a single cause for global amphibian declines and present a three-tiered hierarchical approach that addresses interactions among and between ultimate and proximate factors that contribute to amphibian declines. There are two immediate (proximate) causes of amphibian declines: death and decreased recruitment (reproductive failure). Although much attention has focused on death, few studies have addressed factors that contribute to declines as a result of failed recruitment. Further, a great deal of attention has focused on the role of pathogens in inducing diseases that cause death, but we suggest that pathogen success is profoundly affected by four other ultimate factors: atmospheric change, environmental pollutants, habitat modification and invasive species. Environmental pollutants arise as likely important factors in amphibian declines because they have realized potential to affect recruitment. Further, many studies have documented immunosuppressive effects of pesticides, suggesting a role for environmental contaminants in increased pathogen virulence and disease rates. Increased attention to recruitment and ultimate factors that interact with pathogens is important in addressing this global crisis.

Understanding the vertebrate immune system: insights from the reptilian perspective

Abstract: Reptiles are ectothermic amniotes, providing the key link between ectothermic anamniotic fishes and amphibians, and endothermic amniotic birds and mammals. A greater understanding of reptilian immunity will provide important insights into the evolutionary history of vertebrate immunity as well as the growing field of eco-immunology. Like mammals, reptile immunity is complex and involves innate, cell-mediated and humoral compartments but, overall, there is considerably less known about immune function in reptiles. We review the current literature on each branch of the reptilian immune system, placing this information in context to other vertebrates. Further, we identify key areas that are prime for research as well as areas that are lagging because of lack of reagents in non-model systems.

Phenotypic plasticity and genetic adaptation to high-altitude hypoxia in vertebrates

Abstract: High-altitude environments provide ideal testing grounds for investigations of mechanism and process in physiological adaptation. In vertebrates, much of our understanding of the acclimatization response to high-altitude hypoxia derives from studies of animal species that are native to lowland environments. Such studies can indicate whether phenotypic plasticity will generally facilitate or impede adaptation to high altitude. Here, we review general mechanisms of physiological acclimatization and genetic adaptation to high-altitude hypoxia in birds and mammals. We evaluate whether the acclimatization response to environmental hypoxia can be regarded generally as a mechanism of adaptive phenotypic plasticity, or whether it might sometimes represent a misdirected response that acts as a hindrance to genetic adaptation. In cases in which the acclimatization response to hypoxia is maladaptive, selection will favor an attenuation of the induced phenotypic change. This can result in a form of cryptic adaptive evolution in which phenotypic similarity between high- and low-altitude populations is attributable to directional selection on genetically based trait variation that offsets environmentally induced changes. The blunted erythropoietic and pulmonary vasoconstriction responses to hypoxia in Tibetan humans and numerous high-altitude birds and mammals provide possible examples of this phenomenon. When lowland animals colonize high-altitude environments, adaptive phenotypic plasticity can mitigate the costs of selection, thereby enhancing prospects for population establishment and persistence. By contrast, maladaptive plasticity has the opposite effect. Thus, insights into the acclimatization response of lowland animals to high-altitude hypoxia can provide a basis for predicting how altitudinal range limits might shift in response to climate change.

Physiological climatic limits in Drosophila: patterns and implications.

Abstract: Physiological limits determine susceptibility to environmental changes, and can be assessed at the individual, population or species/lineage levels. Here I discuss these levels in Drosophila, and consider implications for determining species susceptibility to climate change. Limits at the individual level in Drosophila depend on experimental technique and on the context in which traits are evaluated. At the population level, evidence from selection experiments particularly involving Drosophila melanogaster indicate high levels of heritable variation and evolvability for coping with thermal stresses and aridity. An exception is resistance to high temperatures, which reaches a plateau in selection experiments and has a low heritability/evolvability when temperatures are ramped up to a stressful level. In tropical Drosophila species, populations are limited in their ability to evolve increased desiccation and cold resistance. Population limits can arise from trait and gene interactions but results from different laboratory studies are inconsistent and likely to underestimate the strength of interactions under field conditions. Species and lineage comparisons suggest phylogenetic conservatism for resistance to thermal extremes and other stresses. Plastic responses set individual limits but appear to evolve slowly in Drosophila. There is more species-level variation in lower thermal limits and desiccation resistance compared with upper limits, which might reflect different selection pressures and/or low evolvability. When extremes are considered, tropical Drosophila species do not appear more threatened than temperate species by higher temperatures associated with global warming, contrary to recent conjectures. However, species from the humid tropics may be threatened if they cannot adapt genetically to drier conditions.

Epigenetics and transgenerational transfer: A physiological perspective

Abstract: Epigenetics, the transgenerational transfer of phenotypic characters without modification of gene sequence, is a burgeoning area of study in many disciplines of biology. However, the potential impact of this phenomenon on the physiology of animals is not yet broadly appreciated, in part because the phenomenon of epigenetics is not typically part of the design of physiological investigations. Still enigmatic and somewhat ill defined is the relationship between the overarching concept of epigenetics and interesting transgenerational phenomena (e.g. 'maternal/parental effects') that alter the physiological phenotype of subsequent generations. The lingering effect on subsequent generations of an initial environmental disturbance in parent animals can be profound, with genes continuing to be variously silenced or expressed without an associated change in gene sequence for many generations. Known epigenetic mechanisms involved in this phenomenon include chromatin remodeling (DNA methylation and histone modification), RNA-mediated modifications (non-coding RNA and microRNA), as well as other less well studied mechanisms such as self-sustaining loops and structural inheritance. In this review we: (1) discuss how the concepts of epigenetics and maternal effects both overlap with, and are distinct from, each other; (2) analyze examples of existing animal physiological studies based on these concepts; and (3) offer a construct by which to integrate these concepts into the design of future investigations in animal physiology.

Diversity, decoys and the dilution effect: how ecological communities affect disease risk

Abstract: Growing interest in ecology has recently focused on the hypothesis that community diversity can mediate infection levels and disease ('dilution effect'). In turn, biodiversity loss--a widespread consequence of environmental change--can indirectly promote increases in disease, including those of medical and veterinary importance. While this work has focused primarily on correlational studies involving vector-borne microparasite diseases (e.g. Lyme disease, West Nile virus), we argue that parasites with complex life cycles (e.g. helminths, protists, myxosporeans and many fungi) offer an excellent additional model in which to experimentally address mechanistic questions underlying the dilution effect. Here, we unite recent ecological research on the dilution effect in microparasites with decades of parasitological research on the decoy effect in macroparasites to explore key questions surrounding the relationship between community structure and disease. We find consistent evidence that community diversity significantly alters parasite transmission and pathology under laboratory as well as natural conditions. Empirical examples and simple transmission models highlight the diversity of mechanisms through which such changes occur, typically involving predators, parasite decoys, low competency hosts or other parasites. However, the degree of transmission reduction varies among diluting species, parasite stage, and across spatial scales, challenging efforts to make quantitative, taxon-specific predictions about disease. Taken together, this synthesis highlights the broad link between community structure and disease while underscoring the importance of mitigating ongoing changes in biological communities owing to species introductions and extirpations.

The proteomic response of the mussel congeners Mytilus galloprovincialis and M. trossulus to acute heat stress: implications for thermal tolerance limits and metabolic costs of thermal stress.

Abstract: The Mediterranean blue mussel, Mytilus galloprovincialis , an invasive species in California, has displaced the more heat-sensitive native congener, Mytilus trossulus , from its former southern range, possibly due to climate change. By comparing the response of their proteomes to acute heat stress we sought to identify responses common to both species as well as differences that account for greater heat tolerance in the invasive. Mussels were acclimated to 13°C for four weeks and exposed to acute heat stress (24°C, 28°C and 32°C) for 1 h and returned to 13°C to recover for 24 h. Using two-dimensional gel electrophoresis and tandem mass spectrometry we identified 47 and 61 distinct proteins that changed abundance in M. galloprovincialis and M. trossulus , respectively. The onset temperatures of greater abundance of some members of the heat shock protein (Hsp) 70 and small Hsp families were lower in M. trossulus . The abundance of proteasome subunits was lower in M. galloprovincialis but greater in M. trossulus in response to heat. Levels of several NADH-metabolizing proteins, possibly linked to the generation of reactive oxygen species (ROS), were lower at 32°C in the cold-adapted M. trossulus whereas proteins generating NADPH, important in ROS defense, were higher in both species. The abundance of oxidative stress proteins was lower at 32°C in M. trossulus only, indicating that its ability to combat heat-induced oxidative stress is limited to lower temperatures. Levels of NAD-dependent deacetylase (sirtuin 5), which are correlated with lifespan, were lower in M. trossulus in response to heat stress. In summary, the expression patterns of proteins involved in molecular chaperoning, proteolysis, energy metabolism, oxidative damage, cytoskeleton and deacetylation revealed a common loci of heat stress in both mussels but also showed a lower sensitivity to high-temperature damage in the warm-adapted M. galloprovincialis , which is consistent with its expanding range in warmer waters. * AAT : aspartate aminotransferase cdc42 : cell division control protein homolog 42 cMDH : cytosolic malate dehydrogenase CoA : coenzyme A EGFR : epidermal growth factor EST : expressed sequence tag ETC : electron transport chain Hsp : heat shock protein IDH : isocitrate dehydrogenase IPG : immobilized pH gradient MAPK : mitogen-activated protein kinase mMDH : mitochondrial malate dehydrogenase MOWSE : molecular weight search MS : mass spectrometry MVP : major vault protein NAD(H) : nicotinamide adenine dinucleotide (reduced form) NADP(H) : nicotinamide adenine dinucleotide phosphate (reduced form) PCA : principle component analysis PDH : pyruvate dehydrogenase PEPCK : phosphoenolpyruvate carboxykinase pI : isoelectric point PMF : peptide mass fingerprint PTM : post-translational modification ROS : reactive oxygen species sHsps : small heat shock proteins SOD : superoxide dismutase TFA : trifluoroacetic acid T on : onset temperature 2-D GE : two-dimensional gel electrophoresis

Challenges in predicting climate and environmental effects on vector-borne disease episystems in a changing world.

Abstract: Vector-borne pathogens cause enormous suffering to humans and animals. Many are expanding their range into new areas. Dengue, West Nile and Chikungunya have recently caused substantial human epidemics. Arthropod-borne animal diseases like Bluetongue, Rift Valley fever and African horse sickness pose substantial threats to livestock economies around the world. Climate change can impact the vector-borne disease epidemiology. Changes in climate will influence arthropod vectors, their life cycles and life histories, resulting in changes in both vector and pathogen distribution and changes in the ability of arthropods to transmit pathogens. Climate can affect the way pathogens interact with both the arthropod vector and the human or animal host. Predicting and mitigating the effects of future changes in the environment like climate change on the complex arthropod-pathogen-host epidemiological cycle requires understanding of a variety of complex mechanisms from the molecular to the population level. Although there has been substantial progress on many fronts the challenges to effectively understand and mitigate the impact of potential changes in the environment on vector-borne pathogens are formidable and at an early stage of development. The challenges will be explored using several arthropod-borne pathogen systems as illustration, and potential avenues to meet the challenges will be presented.

High-precision tracking of sperm swimming fine structure provides strong test of resistive force theory

Abstract: The shape of the flagellar beat determines the path along which a sperm cell swims. If the flagellum bends periodically about a curved mean shape then the sperm will follow a path with non-zero curvature. To test a simple hydrodynamic theory of flagellar propulsion known as resistive force theory, we conducted high-precision measurements of the head and flagellum motions during circular swimming of bull spermatozoa near a surface. We found that the fine structure of sperm swimming represented by the rapid wiggling of the sperm head around an averaged path is, to high accuracy, accounted for by resistive force theory and results from balancing forces and torques generated by the beating flagellum. We determined the anisotropy ratio between the normal and tangential hydrodynamic friction coefficients of the flagellum to be 1.81+/-0.07 (mean+/-s.d.). On time scales longer than the flagellar beat cycle, sperm cells followed circular paths of non-zero curvature. Our data show that path curvature is approximately equal to twice the average curvature of the flagellum, consistent with quantitative predictions of resistive force theory. Hence, this theory accurately predicts the complex trajectories of sperm cells from the detailed shape of their flagellar beat across different time scales.

On the role of form and kinematics on the hydrodynamics of self-propelled body/caudal fin swimming

Abstract: We carry out fluid-structure interaction simulations of self-propelled virtual swimmers to investigate the effects of body shape (form) and kinematics on the hydrodynamics of undulatory swimming. To separate the effects of form and kinematics, we employ four different virtual swimmers: a carangiform swimmer (i.e. a mackerel swimming like mackerel do in nature); an anguilliform swimmer (i.e. a lamprey swimming like lampreys do in nature); a hybrid swimmer with anguilliform kinematics but carangiform body shape (a mackerel swimming like a lamprey); and another hybrid swimmer with carangiform kinematics but anguilliform body shape (a lamprey swimming like a mackerel). By comparing the performance of swimmers with different kinematics but similar body shapes we study the effects of kinematics whereas by comparing swimmers with similar kinematics but different body shapes we study the effects of form. We show that the anguilliform kinematics not only reaches higher velocities but is also more efficient in the viscous (Re approximately 10(2)) and transitional (Re approximately 10(3)) regimes. However, in the inertial regime (Re=infinity) carangiform kinematics achieves higher velocities and is also more efficient than the anguilliform kinematics. The mackerel body achieves higher swimming speeds in all cases but is more efficient in the inertial regime only whereas the lamprey body is more efficient in the transitional regime. We also show that form and kinematics have little overall effect on the 3-D structure of the wake (i.e. single vs double row vortex streets), which mainly depends on the Strouhal number. Nevertheless, body shape is found to somewhat affect the small-scale features and complexity of the vortex rings shed by the various swimmers.

Balance responses to lateral perturbations in human treadmill walking

Abstract: During walking on a treadmill 10 human subjects (mean age 20 years) were perturbed by 100 ms pushes or pulls to the left or the right, of various magnitudes and in various phases of the gait cycle. Balance was maintained by (1) a stepping strategy (synergy), in which the foot at the next step is positioned a fixed distance outward of the 'extrapolated centre of mass', and (2) a lateral ankle strategy, which comprises a medial or lateral movement of the centre of pressure under the foot sole. The extrapolated centre of mass is defined as the centre of mass position plus the centre of mass velocity multiplied by a parameter related to the subject's leg length. The ankle strategy is the fastest, with a mechanical delay of about 200 ms (20% of a stride), but it can displace the centre of pressure no more than 2 cm. The stepping strategy needs at least 300 ms (30% of a stride) before foot placement, but has a range of 20 cm. When reaction time is sufficient, the magnitude of the total response is in good agreement with our hypothesis: mean centre of pressure (foot) position is a constant distance outward of the extrapolated centre of mass. If the reaction time falls short, a further correction is applied in the next step. In the healthy subjects studied here, no further corrections were necessary, so balance was recovered within two steps (one stride).

Transcriptomic responses to heat stress in invasive and native blue mussels (genus Mytilus): molecular correlates of invasive success.

Abstract: SUMMARY Invasive species are increasingly prevalent in marine ecosystems worldwide. Although many studies have examined the ecological effects of invasives, little is known about the physiological mechanisms that might contribute to invasive success. The mussel Mytilus galloprovincialis , a native of the Mediterranean Sea, is a successful invader on the central and southern coasts of California, where it has largely displaced the native congener, Mytilus trossulus . It has been previously shown that thermal responses of several physiological traits may underlie the capacity of M. galloprovincialis to out-compete M. trossulus in warm habitats. To elucidate possible differences in stress-induced gene expression between these congeners, we developed an oligonucleotide microarray with 8874 probes representing 4488 different genes that recognized mRNAs of both species. In acute heat-stress experiments, 1531 of these genes showed temperature-dependent changes in expression that were highly similar in the two congeners. By contrast, 96 genes showed species-specific responses to heat stress, functionally characterized by their involvement in oxidative stress, proteolysis, energy metabolism, ion transport, cell signaling and cytoskeletal reorganization. The gene that showed the biggest difference between the species was the gene for the molecular chaperone small heat shock protein 24, which was highly induced in M. galloprovincialis and showed only a small change in M. trossulus . These different responses to acute heat stress may help to explain – and predict – the invasive success of M. galloprovincialis in a warming world.

The effects of arm swing on human gait stability

Abstract: Arm swing during human gait has been shown to reduce both angular momentum about the vertical and energy expenditure, and has been hypothesized to enhance gait stability. To examine this hypothesis, we studied the effect of arm swing on the local and global stability of steady-state gait, as well as the ability to perform adequate recovery actions following a perturbation. Trunk kinematics of 11 male subjects was measured in treadmill walking with normal and with restricted arm swing. In half of the trials, gait was perturbed by a position-controlled forward pull to the trunk. We constructed state spaces using data recorded from the unperturbed steady-state walking trials, and quantified local gait stability by calculating maximum Lyapunov exponents. In addition, we analyzed perturbation forces, the distance from the unperturbed gait pattern, and the return toward the normal gait pattern following an external perturbation. Walking without arm swing led to a non-significantly lower Lyapunov exponent (P=0.06), significantly higher perturbation forces (P<0.05), and significantly slower movements away from the attractor (P<0.01). These results suggest that gait without arm swing is characterized by similar local stability to gait with arm swing and a higher perturbation resistance. However, return towards the normal gait pattern was significantly slower (P<0.05) when walking with restricted arms, suggesting that the arms play an important role in the recovery from a perturbation. Collectively, the results suggest that arm swing as such does not enhance gait stability, but rather that recovery movements of the arms contribute to the overall stability of human gait.

Organismal climatology: analyzing environmental variability at scales relevant to physiological stress.

Abstract: Predicting when, where and with what magnitude climate change is likely to affect the fitness, abundance and distribution of organisms and the functioning of ecosystems has emerged as a high priority for scientists and resource managers. However, even in cases where we have detailed knowledge of current species' range boundaries, we often do not understand what, if any, aspects of weather and climate act to set these limits. This shortcoming significantly curtails our capacity to predict potential future range shifts in response to climate change, especially since the factors that set range boundaries under those novel conditions may be different from those that set limits today. We quantitatively examine a nine-year time series of temperature records relevant to the body temperatures of intertidal mussels as measured using biomimetic sensors. Specifically, we explore how a ‘climatology’ of body temperatures, as opposed to long-term records of habitat-level parameters such as air and water temperatures, can be used to extrapolate meaningful spatial and temporal patterns of physiological stress. Using different metrics that correspond to various aspects of physiological stress (seasonal means, cumulative temperature and the return time of extremes) we show that these potential environmental stressors do not always occur in synchrony with one another. Our analysis also shows that patterns of animal temperature are not well correlated with simple, commonly used metrics such as air temperature. Detailed physiological studies can provide guidance to predicting the effects of global climate change on natural ecosystems but only if we concomitantly record, archive and model environmental signals at appropriate scales.

Predicting the impacts of climate change on a globally distributed species: the case of the loggerhead turtle

Abstract: Marine turtles utilise terrestrial and marine habitats and several aspects of their life history are tied to environmental features that are altering due to rapid climate change. We overview the likely impacts of climate change on the biology of these species, which are likely centred upon the thermal ecology of this taxonomic group. Then, focusing in detail on three decades of research on the loggerhead turtle ( Caretta caretta L.), we describe how much progress has been made to date and how future experimental and ecological focus should be directed. Key questions include: what are the current hatchling sex ratios from which to measure future climate-induced changes? What are wild adult sex ratios and how many males are necessary to maintain a fertile and productive population? How will climate change affect turtles in terms of their distribution?

The effects of turbulent eddies on the stability and critical swimming speed of creek chub (Semotilus atromaculatus).

Abstract: SUMMARY The effect of turbulent eddy diameter, vorticity and orientation on the 2 min critical swimming speed and stability of creek chub ( Semotilus atromaculatus ) is reported. Turbulent eddies were visualized and their properties were quantified using particle image velocimetry (PIV). Flow fields with an increasing range in eddy diameter were created by inserting cylinder arrays upstream from the swimming test section. Eddy vorticity increased with increasing velocity. Two orientations of eddies, eddies spinning about a vertical axis and eddies spinning about a horizontal (wall-to-wall) axis, were investigated. Stability challenges were not observed until the largest (95th percentile) eddy diameters reached 76% of the fish body total length. Under these conditions fish were observed to spin in an orientation consistent with the rotational axis of the large eddies and translate downstream. These losses in postural control were termed ‘spills’. Spills were 230% more frequent and lasted 24% longer in turbulent flow fields dominated by horizontal eddies than by vertical eddies of the same diameter. The onset of spills coincided with a 10% and 22% reduction in critical swimming speed in turbulent flows dominated by large vertical and horizontal eddies, respectively. These observations confirm predictions by Pavlov et al., Cada and Odeh, Lupandin, and Liao that the eddy diameter, vorticity and orientation play an important role in the swimming capacity of fishes.

Harbor seal vibrissa morphology suppresses vortex-induced vibrations.

Abstract: SUMMARY Harbor seals ( Phoca vitulina ) often live in dark and turbid waters, where their mystacial vibrissae, or whiskers, play an important role in orientation. Besides detecting and discriminating objects by direct touch, harbor seals use their whiskers to analyze water movements, for example those generated by prey fish or by conspecifics. Even the weak water movements left behind by objects that have passed by earlier can be sensed and followed accurately (hydrodynamic trail following). While scanning the water for these hydrodynamic signals at a swimming speed in the order of meters per second, the seal keeps its long and flexible whiskers in an abducted position, largely perpendicular to the swimming direction. Remarkably, the whiskers of harbor seals possess a specialized undulated surface structure, the function of which was, up to now, unknown. Here, we show that this structure effectively changes the vortex street behind the whiskers and reduces the vibrations that would otherwise be induced by the shedding of vortices from the whiskers (vortex-induced vibrations). Using force measurements, flow measurements and numerical simulations, we find that the dynamic forces on harbor seal whiskers are, by at least an order of magnitude, lower than those on sea lion ( Zalophus californianus ) whiskers, which do not share the undulated structure. The results are discussed in the light of pinniped sensory biology and potential biomimetic applications.

Human walking isn't all hard work: evidence of soft tissue contributions to energy dissipation and return

Abstract: The muscles and tendons of the lower extremity are generally considered the dominant producers of positive and negative work during gait. However, soft-tissue deformations not captured by joint rotations might also dissipate, store and even return substantial energy to the body. A key locomotion event is the collision of the leg with the ground, which deforms soft tissues appreciably in running. Significant deformation might also result from the impulsive ground collision in walking. In a study of normal human walking (N=10; 0.7-2.0 m s(-1) speeds), we show indirect evidence for both negative and positive work performed by soft tissue, consistent with a damped elastic collision and rebound. We used the difference between measured joint work and another quantity - the work performed on the body center of mass - to indicate possible work performed by soft tissue. At 1.25 m s(-1), we estimated that soft tissue performs approximately 7.5 J of negative work per collision. This constitutes approximately 60% of the total negative collision work and 31% of the total negative work per stride. The amount of soft tissue work during collision increases sharply with speed. Each collision is followed by 4 J of soft tissue rebound that is also not captured by joint work measures. Soft tissue deformation may save muscles the effort of actively dissipating energy, and soft tissue elastic rebound could save up to 14% of the total positive work per stride. Soft tissues not only cushion impacts but also appear to perform substantial work.

Changes in materials properties explain the effects of humidity on gecko adhesion

Abstract: SUMMARY Geckos owe their remarkable stickiness to millions of dry setae on their toes, and the mechanism of adhesion in gecko setae has been the topic of scientific scrutiny for over two centuries. Previously, we demonstrated that van der Waals forces are sufficient for strong adhesion and friction in gecko setae, and that water-based capillary adhesion is not required. However, recent studies demonstrated that adhesion increases with relative humidity (RH) and proposed that surface hydration and capillary water bridge formation is important or even necessary. In this study, we confirmed a significant effect of RH on gecko adhesion, but rejected the capillary adhesion hypothesis. While contact forces of isolated tokay gecko setal arrays increased with humidity, the increase was similar on hydrophobic and hydrophilic surfaces, inconsistent with a capillary mechanism. Contact forces increased with RH even at high shear rates, where capillary bridge formation is too slow to affect adhesion. How then can a humidity-related increase in adhesion and friction be explained? The effect of RH on the mechanical properties of setal β-keratin has escaped consideration until now. We discovered that an increase in RH softens setae and increases viscoelastic damping, which increases adhesion. Changes in setal materials properties, not capillary forces, fully explain humidity-enhanced adhesion, and van der Waals forces remain the only empirically supported mechanism of adhesion in geckos.

Aquaporin expression dynamics in osmoregulatory tissues of Atlantic salmon during smoltification and seawater acclimation

Abstract: Osmotic balance in fish is maintained through the coordinated regulation of water and ion transport performed by epithelia in intestine, kidney and gill. In the current study, six aquaporin (AQP) isoforms found in Atlantic salmon (Salmo salar) were classified and their tissue specificity and mRNA expression in response to a hyperosmotic challenge and during smoltification were examined. While AQP-1a was generic, AQP-1b had highest expression in kidney and AQP-3 was predominantly found in oesophagus, gill and muscle. Two novel teleost isoforms, AQP-8a and -8b, were expressed specifically in liver and intestinal segments, respectively. AQP-10 was predominantly expressed in intestinal segments, albeit at very low levels. Transfer from freshwater (FW) to seawater (SW) induced elevated levels of intestinal AQP-1a, -1b and -8b mRNA, whereas only AQP-8b was stimulated during smoltification. In kidney, AQP-1a, -3 and -10 were elevated in SW whereas AQP-1b was reduced compared with FW levels. Correspondingly, renal AQP-1a and -10 peaked during smoltification in April and March, respectively, as AQP-1b and AQP-3 declined. In the gill, AQP-1a and AQP-3 declined in SW whereas AQP-1b increased. Gill AQP-1a and -b peaked in April, whereas AQP-3 declined through smoltification. These reciprocal isoform shifts in renal and gill tissues may be functionally linked with the changed role of these organs in FW compared with SW. The presence and observed dynamics of the AQP-8b isoform specifically in intestinal sections suggest that this is a key water channel responsible for water uptake in the intestinal tract of seawater salmonids.

Slight differences in incubation temperature affect early growth and stress endocrinology of wood duck (Aix sponsa) ducklings.

Abstract: Early developmental experiences, such as incubation conditions, can have important consequences for post-hatching fitness in birds. Although the effects of incubation temperature on phenotype of avian hatchlings are poorly understood, recent research suggests that subtle changes in incubation conditions can influence hatchling characteristics, including body size and condition. We designed an experiment to explore the effects of incubation temperature on hatching success, survival to 9 days post hatch, growth and the hypothalamo-pituitary-adrenal (HPA) axis in wood ducks (Aix sponsa). Wood duck eggs were collected from nest boxes and experimentally incubated at three temperatures (35.0, 35.9 and 37.0 degrees C), each falling within the range of temperatures of naturally incubated wood duck nests. Survival and growth were monitored in ducklings fed ad libitum for 9 days post hatch. In addition, baseline and stress-induced plasma corticosterone concentrations were measured in 2 and 9 day old ducklings. Hatching success and survival to 9 days was greatest in ducks incubated at the intermediate temperature. Ducklings incubated at 35.9 degrees C and 37.0 degrees C had 43% higher growth rates than ducklings incubated at 35.0 degrees C. In addition, ducklings incubated at 35.0 degrees C had higher baseline (17-50%) and stress-induced (32-84%) corticosterone concentrations than ducklings incubated at 35.9 degrees C and 37.0 degrees C at 2 and 9 days post hatch. We also found a significant negative correlation between body size and plasma corticosterone concentrations (baseline and stress-induced) in 9 day old ducklings. To our knowledge, this is the first study to demonstrate that thermal conditions experienced during embryonic development can influence the HPA axis of young birds. Our results illustrate that subtle changes (<1.0 degrees C) in the incubation environment can have important consequences for physiological traits important to fitness.

Sound detection by the longfin squid (Loligo pealeii) studied with auditory evoked potentials: sensitivity to low-frequency particle motion and not pressure

Abstract: SUMMARY Although hearing has been described for many underwater species, there is much debate regarding if and how cephalopods detect sound. Here we quantify the acoustic sensitivity of the longfin squid (Loligo pealeii) using near-field acoustic and shakergenerated acceleration stimuli. Sound field pressure and particle motion components were measured from 30 to 10,000Hz and acceleration stimuli were measured from 20 to 1000Hz. Responses were determined using auditory evoked potentials (AEPs) with electrodes placed near the statocysts. Evoked potentials were generated by both stimuli and consisted of two wave types: (1) rapid stimulus-following waves, and (2) slower, high-amplitude waves, similar to some fish AEPs. Responses were obtained between 30 and 500Hz with lowest thresholds between 100 and 200Hz. At the best frequencies, AEP amplitudes were often >20V. Evoked potentials were extinguished at all frequencies if (1) water temperatures were less than 8°C, (2) statocysts were ablated, or (3) recording electrodes were placed in locations other than near the statocysts. Both the AEP response characteristics and the range of responses suggest that squid detect sound similarly to most fish, with the statocyst acting as an accelerometer through which squid detect the particle motion component of a sound field. The modality and frequency range indicate that squid probably detect acoustic particle motion stimuli from both predators and prey as well as low-frequency environmental sound signatures that may aid navigation. Supplementary material available online at http://jeb.biologists.org/cgi/content/full/213/21/3748/DC1

Immediate spectral flexibility in singing chiffchaffs during experimental exposure to highway noise

Abstract: Sound plays an important role in the life of many animals, including many bird species. Typically, male birds sing to defend a territory and to attract mates. Ambient noise may negatively affect the signal efficiency of their songs, which may be critical to reproductive success. Consequently, anthropogenic noise may be detrimental to individual birds and to populations in cities and along highways. Several bird species that are still common in urban areas have been shown to sing at higher frequency at locations where there is more low-frequency traffic noise. Here we show that chiffchaffs along noisy highways also sing with a higher minimum frequency than chiffchaffs nearby at a quiet riverside. Furthermore, through experimental exposure to highway noise we show that these birds are capable of making such adjustments over a very short time scale. The first 10 songs sung during the noise exposure revealed an immediate shift to higher frequencies, with a return to pre-exposure levels in recordings without noise the following day. In a transmission re-recording experiment we tested the impact of a potential measurement artifact by recording playback of the same songs repeatedly under different controlled noise conditions. We found an upward shift in the minimum frequency measurement associated with more noisy recordings of the same song, but this artifact was not of a scale that it could explain the noise-dependent spectral shifts in chiffchaffs.

What are the physiological and immunological responses of coral to climate warming and disease

Abstract: Coral mortality due to climate-associated stress is likely to increase as the oceans get warmer and more acidic. Coral bleaching and an increase in infectious disease are linked to above average sea surface temperatures. Despite the uncertain future for corals, recent studies have revealed physiological mechanisms that improve coral resilience to the effects of climate change. Some taxa of bleached corals can increase heterotrophic food intake and exchange symbionts for more thermally tolerant clades; this plasticity can increase the probability of surviving lethal thermal stress. Corals can fight invading pathogens with a suite of innate immune responses that slow and even arrest pathogen growth and reduce further tissue damage. Several of these responses, such as the melanin cascade, circulating amoebocytes and antioxidants, are induced in coral hosts during pathogen invasion or disease. Some components of immunity show thermal resilience and are enhanced during temperature stress and even in bleached corals. These examples suggest some plasticity and resilience to cope with environmental change and even the potential for evolution of resistance to disease. However, there is huge variability in responses among coral species, and the rate of climate change is projected to be so rapid that only extremely hardy taxa are likely to survive the projected changes in climate stressors.

Genetic differences in hemoglobin function between highland and lowland deer mice

Abstract: SUMMARY In high-altitude vertebrates, adaptive changes in blood–O 2 affinity may be mediated by modifications of hemoglobin (Hb) structure that affect intrinsic O 2 affinity and/or responsiveness to allosteric effectors that modulate Hb–O 2 affinity. This mode of genotypic specialization is considered typical of mammalian species that are high-altitude natives. Here we investigated genetically based differences in Hb–O 2 affinity between highland and lowland populations of the deer mouse ( Peromyscus maniculatus ), a generalist species that has the broadest altitudinal distribution of any North American mammal. The results of a combined genetic and proteomic analysis revealed that deer mice harbor a high level of Hb isoform diversity that is attributable to allelic polymorphism at two tandemly duplicated α-globin genes and two tandemly duplicated β-globin genes. This high level of isoHb diversity translates into a correspondingly high level of interindividual variation in Hb functional properties. O 2 equilibrium experiments revealed that the Hbs of highland mice exhibit slightly higher intrinsic O 2 affinities and significantly lower Cl – sensitivities relative to the Hbs of lowland mice. The experiments also revealed distinct biochemical properties of deer mouse Hb related to the anion-dependent allosteric regulation of O 2 affinity. In conjunction with previous findings, our results demonstrate that modifications of Hb structure that alter allosteric anion sensitivity play an important role in the adaptive fine-tuning of blood–O 2 affinity.

Individual variation and repeatability in aerobic and anaerobic swimming performance of European sea bass, Dicentrarchus labrax.

Abstract: SUMMARY Studies of inter-individual variation in fish swimming performance may provide insight into how selection has influenced diversity in phenotypic traits. We investigated individual variation and short-term repeatability of individual swimming performance by wild European sea bass in a constant acceleration test (CAT). Fish were challenged with four consecutive CATs with 5 min rest between trials. We measured maximum anaerobic speed at exhaustion ( U CAT ), gait transition speed from steady aerobic to unsteady anaerobic swimming ( U gt ), routine metabolic rate (RMR), post-CAT maximum metabolic rate (MMR), aerobic scope and recovery time from the CATs. Fish achieved significantly higher speeds during the first CAT ( U CAT =170 cm s –1 ), and had much more inter-individual variation in performance (coefficient of variation, CV=18.43%) than in the subsequent three tests ( U CAT =134 cm s –1 ; CV=7.3%), which were very repeatable among individuals. The individual variation in U CAT in the first trial could be accounted for almost exclusively by variation in anaerobic burst-and-coast performance beyond U gt . The U gt itself varied substantially between individuals (CV=11.4%), but was significantly repeatable across all four trials. Individual RMR and MMR varied considerably, but the rank order of post-CAT MMR was highly repeatable. Recovery rate from the four CATs was highly variable and correlated positively with the first U CAT (longer recovery for higher speeds) but negatively with RMR and aerobic scope (shorter recovery for higher RMR and aerobic scope). This large variation in individual performance coupled with the strong correlations between some of the studied variables may reflect divergent selection favouring alternative strategies for foraging and avoiding predation.