Showing papers in "The Journal of Experimental Biology in 2001"

The control of flight force by a flapping wing: lift and drag production.

Abstract: We used a dynamically scaled mechanical model of the fruit fly Drosophila melanogaster to study how changes in wing kinematics influence the production of unsteady aerodynamic forces in insect flight. We examined 191 separate sets of kinematic patterns that differed with respect to stroke amplitude, angle of attack, flip timing, flip duration and the shape and magnitude of stroke deviation. Instantaneous aerodynamic forces were measured using a two-dimensional force sensor mounted at the base of the wing. The influence of unsteady rotational effects was assessed by comparing the time course of measured forces with that of corresponding translational quasi-steady estimates. For each pattern, we also calculated mean stroke-averaged values of the force coefficients and an estimate of profile power. The results of this analysis may be divided into four main points. (i) For a short, symmetrical wing flip, mean lift was optimized by a stroke amplitude of 180° and an angle of attack of 50°. At all stroke amplitudes, mean drag increased monotonically with increasing angle of attack. Translational quasi-steady predictions better matched the measured values at high stroke amplitude than at low stroke amplitude. This discrepancy was due to the increasing importance of rotational mechanisms in kinematic patterns with low stroke amplitude. (ii) For a 180° stroke amplitude and a 45° angle of attack, lift was maximized by short-duration flips occurring just slightly in advance of stroke reversal. Symmetrical rotations produced similarly high performance. Wing rotation that occurred after stroke reversal, however, produced very low mean lift. (iii) The production of aerodynamic forces was sensitive to changes in the magnitude of the wing’s deviation from the mean stroke plane (stroke deviation) as well as to the actual shape of the wing tip trajectory. However, in all examples, stroke deviation lowered aerodynamic performance relative to the no deviation case. This attenuation was due, in part, to a trade-off between lift and a radially directed component of total aerodynamic force. Thus, while we found no evidence that stroke deviation can augment lift, it nevertheless may be used to modulate forces on the two wings. Thus, insects might use such changes in wing kinematics during steering maneuvers to generate appropriate force moments. (iv) While quasi-steady estimates failed to capture the time course of measured lift for nearly all kinematic patterns, they did predict with reasonable accuracy stroke-averaged values for the mean lift coefficient. However, quasi-steady estimates grossly underestimated the magnitude of the mean drag coefficient under all conditions. This discrepancy was due to the contribution of rotational effects that steady-state estimates do not capture. This result suggests that many prior estimates of mechanical power based on wing kinematics may have been grossly underestimated.

Mechanisms of cell survival in hypoxia and hypothermia.

Abstract: Most animals experience some degree of hypoxia and hypothermia during the course of their natural life history either as a consequence of ambient 'exposure' per se or through metabolic, respiratory and/or circulatory insufficiency. A prevailing experimental approach has been to probe tissues from natural models of hypoxia-tolerant and cold-tolerant vertebrates to look for common mechanisms of defence against O(2) lack and hypothermia. The ability to sustain vital cellular functions in severe cases of either condition varies widely amongst the vertebrates. Like humans, the vast majority of mammals are unable to survive prolonged periods of hypothermia or O(2) deprivation owing to irreversible membrane damage and loss of cellular ion homeostasis in vital organs such as the brain and heart. However, numerous hibernating endotherms, neonatal and diving mammals as well as many ectotherms can tolerate prolonged periods that would, in clinical terms, be called asphyxia or deep hypothermia. The key to their survival under such conditions lies in an inherent ability to downregulate their cellular metabolic rate to new hypometabolic steady states in a way that balances the ATP demand and ATP supply pathways.

Functional and structural adaptations of skeletal muscle to microgravity

Abstract: Our purpose is to summarize the major effects of space travel on skeletal muscle with particular emphasis on factors that alter function. The primary deleterious changes are muscle atrophy and the associated decline in peak force and power. Studies on both rats and humans demonstrate a rapid loss of cell mass with microgravity. In rats, a reduction in muscle mass of up to 37% was observed within 1 week. For both species, the antigravity soleus muscle showed greater atrophy than the fast-twitch gastrocnemius. However, in the rat, the slow type I fibers atrophied more than the fast type II fibers, while in humans, the fast type II fibers were at least as susceptible to space-induced atrophy as the slow fiber type. Space flight also resulted in a significant decline in peak force. For example, the maximal voluntary contraction of the human plantar flexor muscles declined by 20-48% following 6 months in space, while a 21% decline in the peak force of the soleus type I fibers was observed after a 17-day shuttle flight. The reduced force can be attributed both to muscle atrophy and to a selective loss of contractile protein. The former was the primary cause because, when force was expressed per cross-sectional area (kNm(-2)), the human fast type II and slow type I fibers of the soleus showed no change and a 4% decrease in force, respectively. Microgravity has been shown to increase the shortening velocity of the plantar flexors. This increase can be attributed both to an elevated maximal shortening velocity (V(0)) of the individual slow and fast fibers and to an increased expression of fibers containing fast myosin. Although the cause of the former is unknown, it might result from the selective loss of the thin filament actin and an associated decline in the internal drag during cross-bridge cycling. Despite the increase in fiber V(0), peak power of the slow type I fiber was reduced following space flight. The decreased power was a direct result of the reduced force caused by the fiber atrophy. In addition to fiber atrophy and the loss of force and power, weightlessness reduces the ability of the slow soleus to oxidize fats and increases the utilization of muscle glycogen, at least in rats. This substrate change leads to an increased rate of fatigue. Finally, with return to the 1g environment of earth, rat studies have shown an increased occurrence of eccentric contraction-induced fiber damage. The damage occurs with re-loading and not in-flight, but the etiology has not been established.

Muscle tissue adaptations to hypoxia

Abstract: This review reports on the effects of hypoxia on human skeletal muscle tissue. It was hypothesized in early reports that chronic hypoxia, as the main physiological stress during exposure to altitude, per se might positively affect muscle oxidative capacity and capillarity. However, it is now established that sustained exposure to severe hypoxia has detrimental effects on muscle structure. Short-term effects on skeletal muscle structure can readily be observed after 2 months of acute exposure of lowlanders to severe hypoxia, e.g. during typical mountaineering expeditions to the Himalayas. The full range of phenotypic malleability of muscle tissue is demonstrated in people living permanently at high altitude (e.g. at La Paz, 3600-4000 m). In addition, there is some evidence for genetic adaptations to hypoxia in high-altitude populations such as Tibetans and Quechuas, who have been exposed to altitudes in excess of 3500 m for thousands of generations. The hallmark of muscle adaptation to hypoxia in all these cases is a decrease in muscle oxidative capacity concomitant with a decrease in aerobic work capacity. It is thought that local tissue hypoxia is an important adaptive stress for muscle tissue in exercise training, so these results seem contra-intuitive. Studies have therefore been conducted in which subjects were exposed to hypoxia only during exercise sessions. In this situation, the potentially negative effects of permanent hypoxic exposure and other confounding variables related to exposure to high altitude could be avoided. Training in hypoxia results, at the molecular level, in an upregulation of the regulatory subunit of hypoxia-inducible factor-1 (HIF-1). Possibly as a consequence of this upregulation of HIF-1, the levels mRNAs for myoglobin, for vascular endothelial growth factor and for glycolytic enzymes, such as phosphofructokinase, together with mitochondrial and capillary densities, increased in a hypoxia-dependent manner. Functional analyses revealed positive effects on V(O(2)max) (when measured at altitude) on maximal power output and on lean body mass. In addition to the positive effects of hypoxia training on athletic performance, there is some recent indication that hypoxia training has a positive effect on the risk factors for cardiovascular disease.

Recovery periods restore mechanosensitivity to dynamically loaded bone

Abstract: Bone cells are capable of sensing and responding to mechanical forces, but mechanosensitivity begins to decline soon after the stimulus is initiated. Under continued stimulation, bone is desensitized to mechanical stimuli. We sought to determine the amount of time required to restore mechanosensitivity to desensitized bone cells in vivo by manipulating the recovery time (0, 0.5, 1, 2, 4 or 8 h) allowed between four identical daily loading bouts. We also investigated the osteogenic effectiveness of shorter-term recovery periods, lasting several seconds (0.5, 3.5, 7 or 14 s), introduced between each of 36 identical daily loading cycles. Using the rat tibia four-point bending model, the right tibia of 144 adult female Sprague-Dawley rats was subjected to bending, sham bending or no loading. In the rats receiving recovery periods between loading bouts, histomorphometric measurements from the endocortical surface of the loaded and nonloaded control (left) tibiae revealed more than 100 % higher relative bone formation rates in the 8 h recovery group than in the 0 and 0.5 h recovery groups. Approximately 8 h of recovery was sufficient to restore full mechanosensitivity to the cells. In the rats allowed time to recover between load cycles, 14 s of recovery resulted in significantly higher (66-190 %) relative bone formation rates compared to any of the three shorter recovery periods. In both experiments, bone formation in the sham-bending animals was similar to that in the nonloaded control group. The results demonstrate the importance of recovery periods for (i) restoring mechanosensitivity to bone cells and (ii) maximizing the osteogenic effects of mechanical loading (exercise) regimens.

Pectoral fin locomotion in batoid fishes: undulation versus oscillation

Abstract: This study explores the dichotomy between undulatory (passing multiple waves down the fin or body) and oscillatory (flapping) locomotion by comparing the kinematics of pectoral fin locomotion in eight species of batoids (Dasyatis americana, D. sabina, D. say, D. violacea, Gymnura micrura, Raja eglanteria, Rhinobatos lentiginosus and Rhinoptera bonasus) that differ in their swimming behavior, phylogenetic position and lifestyle. The goals of this study are to describe and compare the pectoral fin locomotor behavior of the eight batoid species, to clarify how fin movements change with swimming speed for each species and to analyze critically the undulation/oscillation continuum proposed by Breder using batoids as an example. Kinematic data were recorded for each species over a range of swimming velocities (1-3 disc lengths s(-1)). The eight species in this study vary greatly in their swimming modes. Rhinobatos lentiginosus uses a combination of axial-based and pectoral-fin-based undulation to move forward through the water, with primary thrust generated by the tail. The pectoral fins are activated in short undulatory bursts for increasing swimming speed and for maneuvering. Raja eglanteria uses a combination of pectoral and pelvic locomotion, although only pectoral locomotion is analyzed here. The other six species use pectoral locomotion exclusively to propel themselves through the water. Dasyatis sabina and D. say have the most undulatory fins with an average of 1.3 waves per fin length, whereas Rhinoptera bonasus has the most oscillatory fin behavior with 0.4 waves per fin length. The remaining species range between these two extremes in the degree of undulation present on their fins. There is an apparent trade-off between fin-beat frequency and amplitude. Rhinoptera bonasus has the lowest frequency and the highest fin amplitude, whereas Rhinobatos lentiginosus has the highest frequency and the lowest amplitude among the eight species examined. The kinematic variables that batoids modify to change swimming velocity vary among different species. Rhinobatos lentiginosus increases its tail-beat frequency to increase swimming speed. In contrast, the four Dasyatis species increase swimming speed by increasing frequency and wavespeed, although D. americana also changes wave number. Raja eglanteria modifies its swimming velocity by changing wavespeed and wave number. Rhinoptera bonasus increases wavespeed, Gymnura micrura decreases wave number, and both Rhinoptera bonasus and Gymnura micrura increase fin-tip velocity to increase swimming velocity. Batoid species fall onto a continuum between undulation and oscillation on the basis of the number of waves present on the fins.

Adjusting the thermostat: the threshold induction temperature for the heat-shock response in intertidal mussels (genus Mytilus) changes as a function of thermal history.

Abstract: Spatio-temporal variation in heat-shock gene expression gives organisms the ability to respond to changing thermal environments. The temperature at which heat-shock genes are induced, the threshold induction temperature, varies as a function of the recent thermal history of an organism. To elucidate the mechanism by which this plasticity in gene expression is achieved, we determined heat-shock protein (Hsp) induction threshold temperatures in the intertidal mussel Mytilus trossulus collected from the field in February and again in August. In a separate experiment, threshold induction temperatures, endogenous levels of both the constitutive and inducible isoforms of Hsps from the 70 kDa family and the quantity of ubiquitinated proteins (a measure of cellular protein denaturation) were measured in M. trossulus after either 6 weeks of cold acclimation in the laboratory or acclimatization to warm, summer temperatures in the field over the same period. In addition, we quantified levels of activated heat-shock transcription factor 1 (HSF1) in both groups of mussels (HSF1 inducibly transactivates all classes of Hsp genes). Lastly, we compared the temperature of HSF1 activation with the induction threshold temperature in the congeneric M. californianus. It was found that the threshold induction temperature in M. trossulus was 23 degrees C in February and 28 degrees C in August. This agreed with the acclimation/acclimatization experiment, in which mussels acclimated in seawater tables to a constant temperature of 10-11 degrees C for 6 weeks displayed a threshold induction temperature of 20-23 degrees C compared with 26-29 degrees C for individuals that were experiencing considerably warmer body temperatures in the intertidal zone over the same period. This coincided with a significant increase in the inducible isoform of Hsp70 in warm-acclimatized individuals but no increase in the constitutive isoform or in HSF1. Levels of ubiquitin-conjugated protein were significantly higher in the field mussels than in the laboratory-acclimated individuals. Finally, the temperature of HSF1 activation in M. californianus was found to be approximately 9 degrees C lower than the induction threshold for this species.

Sarcomere length operating range of vertebrate muscles during movement

Abstract: The force generated by skeletal muscle varies with sarcomere length and velocity. An understanding of the sarcomere length changes that occur during movement provides insights into the physiological importance of this relationship and may provide insights into the design of certain muscle/joint combinations. The purpose of this review is to summarize and analyze the available literature regarding published sarcomere length operating ranges reported for various species. Our secondary purpose is to apply analytical techniques to determine whether generalizations can be made regarding the "normal" sarcomere length operating range of skeletal muscle. The analysis suggests that many muscles operate over a narrow range of sarcomere lengths, covering 94+/-13 % of optimal sarcomere length. Sarcomere length measurements are found to be systematically influenced by the rigor state and methods used to make these measurements.

Rapid cold-hardening of Drosophila melanogaster (Diptera: Drosophiladae) during ecologically based thermoperiodic cycles.

Abstract: In contrast to most studies of rapid cold-hardening, in which abrupt transfers to low temperatures are used to induce an acclimatory response, the primary objectives of this study were to determine (i) whether rapid cold-hardening was induced during the cooling phase of an ecologically based thermoperiod, (ii) whether the protection afforded was lost during warming or contributed to increased cold-tolerance during subsequent cycles and (iii) whether the major thermally inducible stress protein (Hsp70) or carbohydrate cryoprotectants contributed to the protection afforded by rapid cold-hardening. During the cooling phase of a single ecologically based thermoperiod, the tolerance of Drosophila melanogaster to 1 h at −7 degrees C increased from 5 +/− 5% survival to 62.5 +/− 7.3% (means +/− S.E.M., N=40-60), while their critical thermal minima (CTmin) decreased by 1.9 degrees C. Cold hardiness increased with the number of thermoperiods to which flies were exposed; i.e. flies exposed to six thermoperiods were more cold-tolerant than those exposed to two. Endogenous levels of Hsp70 and carbohydrate cryoprotectants were unchanged in rapidly cold-hardened adults compared with controls held at a constant 23 degrees C. In nature, rapid cold-hardening probably affords subtle benefits during short-term cooling, such as allowing D. melanogaster to remain active at lower temperatures than they otherwise could.

Evidence that a central governor regulates exercise performance during acute hypoxia and hyperoxia

Abstract: An enduring hypothesis in exercise physiology holds that a limiting cardiorespiratory function determines maximal exercise performance as a result of specific metabolic changes in the exercising skeletal muscle, so-called peripheral fatigue. The origins of this classical hypothesis can be traced to work undertaken by Nobel Laureate A. V. Hill and his colleagues in London between 1923 and 1925. According to their classical model, peripheral fatigue occurs only after the onset of heart fatigue or failure. Thus, correctly interpreted, the Hill hypothesis predicts that it is the heart, not the skeletal muscle, that is at risk of anaerobiosis or ischaemia during maximal exercise. To prevent myocardial damage during maximal exercise, Hill proposed the existence of a ‘governor’ in either the heart or brain to limit heart work when myocardial ischaemia developed. Cardiorespiratory function during maximal exercise at different altitudes or at different oxygen fractions of inspired air provides a definitive test for the presence of a governor and its function. If skeletal muscle anaerobiosis is the protected variable then, under conditions in which arterial oxygen content is reduced, maximal exercise should terminate with peak cardiovascular function to ensure maximum delivery of oxygen to the active muscle. In contrast, if the function of the heart or some other oxygen-sensitive organ is to be protected, then peak cardiovascular function will be higher during hyperoxia and reduced during hypoxia compared with normoxia. This paper reviews the evidence that peak cardiovascular function is reduced during maximal exercise in both acute and chronic hypoxia with no evidence for any primary alterations in myocardial function. Since peak skeletal muscle electromyographic activity is also reduced during hypoxia, these data support a model in which a central, neural governor constrains the cardiac output by regulating the mass of skeletal muscle that can be activated during maximal exercise in both acute and chronic hypoxia.

The metabolic cost of birdsong production.

Abstract: The metabolic cost of birdsong production has not been studied in detail but is of importance in our understanding of how selective pressures shape song behavior. We measured rates of oxygen consumption during song in three songbird species, zebra finches (Taeniopygia guttata), Waterslager canaries (Serinus canaria) and European starlings (Sturnus vulgaris). These species sing songs with different acoustic and temporal characteristics: short stereotyped song (zebra finch), long song with high temporal complexity (canary) and long song with high acoustic, but low temporal, complexity (starling). In all three species, song slightly increased the rate of oxygen consumption over pre-song levels (1.02-1.36-fold). In zebra finches, the metabolic cost per song motif averaged 1.2 microl g(-1). This cost per motif did not change over the range of song duration measured for the four individuals. Surprisingly, the metabolic cost of song production in the species with the temporally most complex song, the canary, was no greater than in the other two species. In starlings, a 16 dB increase in sound intensity was accompanied by a 1.16-fold increase in the rate of oxygen consumption. These data indicate that the metabolic cost of song production in the songbird species studied is no higher than that for other types of vocal behavior in various bird groups. Our analysis shows that the metabolic cost of singing is also similar to that of calling in frogs and of human speech production. However, difficulties with measurements on freely behaving birds in a small respirometry chamber limit the depth of analysis that is possible.

Plasticity of chemotaxis revealed by paired presentation of a chemoattractant and starvation in the nematode Caenorhabditis elegans

Abstract: While the basic functioning of the nervous system of Caenorhabditis elegans has been extensively studied, its behavioural plasticities have not been fully explored because of the limited availability of assay systems. We report here a simple form of chemotaxis plasticity in this organism: when worms are starved on plates that contain NaCl, their chemotaxis towards NaCl falls dramatically. This conditioning requires both the presence of NaCl and the absence of a bacterial food source, indicating that it is not merely adaptation or habituation, but that it is likely to be a form of associative learning. While chemotaxis towards volatile chemoattractants does not change significantly after conditioning with NaCl, chemotaxis towards other water-soluble attractants does decrease. This suggests that an altered response of a cell or a group of cells specifically involved in chemotaxis towards water-soluble chemoattractants is responsible for the behavioural alteration. The decrease in chemotaxis occurred slowly over 3-4 h of conditioning and returned quickly to the original level when either of the conditioning stimuli, NaCl or starvation, was removed. The application of serotonin partially blocked this reduction in chemotaxis, consistent with the proposed function of this neurotransmitter in food signalling. Using this assay, we have isolated three mutants with reduced plasticity. This assay system expands the opportunities for studying the molecular and cellular mechanisms of behavioural plasticity in C. elegans.

Locomotor function of the dorsal fin in teleost fishes: experimental analysis of wake forces in sunfish

Abstract: A key evolutionary transformation of the locomotor system of ray-finned fishes is the morphological elaboration of the dorsal fin. Within Teleostei, the dorsal fin primitively is a single midline structure supported by soft, flexible fin rays. In its derived condition, the fin is made up of two anatomically distinct portions: an anterior section supported by spines, and a posterior section that is soft-rayed. We have a very limited understanding of the functional significance of this evolutionary variation in dorsal fin design. To initiate empirical hydrodynamic study of dorsal fin function in teleost fishes, we analyzed the wake created by the soft dorsal fin of bluegill sunfish (Lepomis macrochirus) during both steady swimming and unsteady turning maneuvers. Digital particle image velocimetry was used to visualize wake structures and to calculate in vivo locomotor forces. Study of the vortices generated simultaneously by the soft dorsal and caudal fins during locomotion allowed experimental characterization of median-fin wake interactions. During high-speed swimming (i.e. above the gait transition from pectoral- to median-fin locomotion), the soft dorsal fin undergoes regular oscillatory motion which, in comparison with analogous movement by the tail, is phase-advanced (by 30% of the cycle period) and of lower sweep amplitude (by 1.0 cm). Undulations of the soft dorsal fin during steady swimming at 1.1 bodylength s(-1) generate a reverse von Karman vortex street wake that contributes 12% of total thrust. During low-speed turns, the soft dorsal fin produces discrete pairs of counterrotating vortices with a central region of high-velocity jet flow. This vortex wake, generated in the latter stage of the turn and posterior to the center of mass of the body, counteracts torque generated earlier in the turn by the anteriorly positioned pectoral fins and thereby corrects the heading of the fish as it begins to translate forward away from the turning stimulus. One-third of the laterally directed fluid force measured during turning is developed by the soft dorsal fin. For steady swimming, we present empirical evidence that vortex structures generated by the soft dorsal fin upstream can constructively interact with those produced by the caudal fin downstream. Reinforcement of circulation around the tail through interception of the dorsal fin's vortices is proposed as a mechanism for augmenting wake energy and enhancing thrust. Swimming in fishes involves the partitioning of locomotor force among several independent fin systems. Coordinated use of the pectoral fins, caudal fin and soft dorsal fin to increase wake momentum, as documented for L. macrochirus, highlights the ability of teleost fishes to employ multiple propulsors simultaneously for controlling complex swimming behaviors.

Cone-based vision of rats for ultraviolet and visible lights

Abstract: Rats (Rattus norvegicus) have two classes of cone, one containing an ultraviolet (UV)-sensitive photopigment and the other housing a pigment maximally sensitive in the middle (M) wavelengths of the visible spectrum. The manner in which signals from these two cone types contribute to rat vision was investigated through recordings of a gross electrical potential (the electroretinogram, ERG) and behavioral discrimination tests. Spectral sensitivity functions obtained from both types of measurement indicate clear contributions from each of the cone classes, but there is a marked enhancement of the relative sensitivity to UV light in the behavioral index; for instance, under some photopic test conditions, rats are approximately equally sensitive to middle-wavelength and UV lights. In adaptation tests, thresholds for UV and M lights were found to be differentially elevated in the presence of chromatic adapting backgrounds, thus providing the possibility that signals from the two cones could be used by the rat visual system to support color discriminations. Evidence of dichromatic color vision in the rat was subsequently obtained from tests of wavelength discrimination.

The boundary layer of swimming fish.

Abstract: Tangential and normal velocity profiles of the boundary layer surrounding live swimming fish were determined by digital particle tracking velocimetry, DPTV. Two species were examined: the scup Stenotomus chrysops, a carangiform swimmer, and the smooth dogfish Mustelus canis, an anguilliform swimmer. Measurements were taken at several locations over the surfaces of the fish and throughout complete undulatory cycles of their propulsive motions. The Reynolds number based on length, Re, ranged from 3x10(3) to 3x10(5). In general, boundary layer profiles were found to match known laminar and turbulent profiles including those of Blasius, Falkner and Skan and the law of the wall. In still water, boundary layer profile shape always suggested laminar flow. In flowing water, boundary layer profile shape suggested laminar flow at lower Reynolds numbers and turbulent flow at the highest Reynolds numbers. In some cases, oscillation between laminar and turbulent profile shapes with body phase was observed. Local friction coefficients, boundary layer thickness and fluid velocities at the edge of the boundary layer were suggestive of local oscillatory and mean streamwise acceleration of the boundary layer. The behavior of these variables differed significantly in the boundary layer over a rigid fish. Total skin friction was determined. Swimming fish were found to experience greater friction drag than the same fish stretched straight in the flow. Nevertheless, the power necessary to overcome friction drag was determined to be within previous experimentally measured power outputs. No separation of the boundary layer was observed around swimming fish, suggesting negligible form drag. Inflected boundary layers, suggestive of incipient separation, were observed sporadically, but appeared to be stabilized at later phases of the undulatory cycle. These phenomena may be evidence of hydrodynamic sensing and response towards the optimization of swimming performance.

Polarization vision--a uniform sensory capacity?

Abstract: In this concept paper, three scenarios are described in which animals make use of polarized light: the underwater world, the water surface and the terrestrial habitat vaulted by the pattern of polarized light in the sky. Within these various visual environments, polarized light is used in a number of ways that make quite different demands on the neural circuitries mediating these different types of behaviour. Apart from some common receptor and pre-processing mechanisms, the underlying neural mechanisms may differ accordingly. Often, information about chi (the angle of polarization), d (the degree of polarization) and lambda (the spectral content) might not --and need not--be disentangled. Hence, the hypothesis entertained in this account is that polarization vision comes in various guises, and that the answer to the question posed in the title is most probably no.

A new technique for monitoring the behaviour of free-ranging Adélie penguins.

Abstract: Measurement of the time allocation of penguins at sea has been a major goal of researchers in recent years. Until now, however, no equipment has been available that would allow measurement of the aquatic and terrestrial behaviour of an Antarctic penguin while it is commuting between the colony and the foraging grounds. A new motion detector, based on the measurement of acceleration, has been used here in addition to current methods of inferring behaviour using data loggers that monitor depth and speed. We present data on the time allocation of Adelie penguins (Pygoscelis adeliae) according to the different types of behaviours they display during their foraging trips: walking, tobogganing, standing on land, lying on land, resting at the water surface, porpoising and diving. To illustrate the potential of this new technique, we compared the behaviour of Adelie penguins during the chick-rearing period in a fast sea-ice region and an ice-free region. The proportion of time spent standing, lying on land and walking during foraging trips was greater for penguins in the sea-ice region (37.6+/-13.3% standing, 21.6+/-15.6% lying and 5.9+/-6.3% walking) than for those in the ice-free region (12.0+/-15.8 % standing, 0.38+/-0.60% lying and 0 % walking), whereas the proportion of time spent resting at the water surface and porpoising was greater for birds in the ice-free region (38.1+/-6.4% resting and 1.1+/-1.1% porpoising) than for those in the sea-ice region (3.0+/-2.3% resting and 0% porpoising; means +/- s.d., N=7 for the sea-ice region, N=4 for the ice-free region). Using this new approach, further studies combining the monitoring of marine resources in different Antarctic sites and the measurement of the energy expenditure of foraging penguins, e.g. using heart rates, will constitute a powerful tool for investigating the effects of environmental conditions on their foraging strategy. This technique will expand our ability to monitor many animals in the field.

Limits to sustained energy intake. III. Effects of concurrent pregnancy and lactation in Mus musculus.

Abstract: To determine whether mice were limited in their capacity to absorb energy during late lactation, we attempted to increase the energy burden experienced by a group of female mice during late lactation by mating them at the postpartum oestrus, hence combining the energy demands of pregnancy and lactation. These experimental mice were therefore concurrently pregnant and lactating in their first lactation, and were followed through a normal second lactation. In a control group, females also underwent two lactations but sequentially, with the second mating after the first litter had been weaned. Maternal mass and food intake were measured throughout the first lactation, second pregnancy and second lactation. Maternal resting metabolic rate (RMR) was measured prior to the first mating and then at the peak of both the first and second lactations. Litter size and litter mass were also measured throughout both lactations. In the first lactation, experimental mice had a lower mass-independent RMR (F1,88=5.15, P=0.026) and raised significantly heavier pups (t=2.77, d.f.=32, P=0.0093) than the control mice. Experimental mice delayed implantation at the start of the second pregnancy. The extent of the delay was positively related to litter size during the first lactation (F1,19=4.58, P=0.046) and negatively related to mean pup mass (F1,19=5.78, P=0.027) in the first lactation. In the second lactation, the experimental mice gave birth to more (t=2.75, d.f.=38, P=0.0092) and lighter (t=-5.01, d.f.=38, P<0.0001) pups than did the controls in their second lactation. Maternal asymptotic daily food intake of control mice in the second lactation was significantly higher (t=-4.39, d.f.=37, P=0.0001) than that of the experimental mice and higher than that of controls during their first lactation. Despite the added burden on the experimental females during their first lactation, there was no increase in their food intake, which suggested that they might be limited by their capacity to absorb energy. However, control females appeared to be capable of increasing their asymptotic food intake beyond the supposed limits estimated previously, suggesting that the previously established limit was not a fixed central limitation on food intake. As RMR increased in parallel with the increase in food intake during the second lactation of control mice, the sustained energy intake remained at around 7.0xRMR.

Frontiers of hypoxia research: acute mountain sickness.

Abstract: Traditionally, scientists and clinicians have explored peripheral physiological responses to acute hypoxia to explain the pathophysiological processes that lead to acute mountain sickness (AMS) and high-altitude cerebral edema (HACE). After more than 100 years of investigation, little is yet known about the fundamental causes of the headache and nausea that are the main symptoms of AMS. Thus, we review the evidence supporting a change in focus to the role of the central nervous system in AMS. Our justification is (i) that the symptoms of AMS and HACE are largely neurological, (ii) that HACE is considered to be the end-stage of severe AMS and was recently identified as a vasogenic edema, opening the door for a role for blood-brain barrier permeability in AMS, (iii) that new, non-invasive techniques make measurement of brain water levels and cerebral blood volume possible and (iv) that the available experimental evidence and theoretical arguments support a significant role for brain swelling in the pathophysiology of AMS. We believe that an examination of the responses of the central nervous system to acute hypoxia will reveal important new pathophysiological processes that may help explain AMS and HACE.

How the clear-sky angle of polarization pattern continues underneath clouds: full-sky measurements and implications for animal orientation

Abstract: One of the biologically most important parameters of the cloudy sky is the proportion P of the celestial polarization pattern available for use in animal navigation. We evaluated this parameter by measuring the polarization patterns of clear and cloudy skies using 180 degrees (full-sky) imaging polarimetry in the red (650 nm), green (550 nm) and blue (450 nm) ranges of the spectrum under clear and partly cloudy conditions. The resulting data were compared with the corresponding celestial polarization patterns calculated using the single-scattering Rayleigh model. We show convincingly that the pattern of the angle of polarization (e-vectors) in a clear sky continues underneath clouds if regions of the clouds and parts of the airspace between the clouds and the earth surface (being shady at the position of the observer) are directly lit by the sun. The scattering and polarization of direct sunlight on the cloud particles and in the air columns underneath the clouds result in the same e-vector pattern as that present in clear sky. This phenomenon can be exploited for animal navigation if the degree of polarization is higher than the perceptual threshold of the visual system, because the angle rather than the degree of polarization is the most important optical cue used in the polarization compass. Hence, the clouds reduce the extent of sky polarization pattern that is useful for animal orientation much less than has hitherto been assumed. We further demonstrate quantitatively that the shorter the wavelength, the greater the proportion of celestial polarization that can be used by animals under cloudy-sky conditions. As has already been suggested by others, this phenomenon may solve the ultraviolet paradox of polarization vision in insects such as hymenopterans and dipterans. The present study extends previous findings by using the technique of 180 degrees imaging polarimetry to measure and analyse celestial polarization patterns.

How the body contributes to the wake in undulatory fish swimming: flow fields of a swimming eel (anguilla anguilla)

Abstract: SUMMARY Undulatory swimmers generate thrust by passing a transverse wave down their body. Thrust is generated not just at the tail, but also to a varying degree by the body, depending on the fish9s morphology and swimming movements. To examine the mechanisms by which the body in particular contributes to thrust production, we chose eels, which have no pronounced tail fin and hence are thought to generate all their thrust with their body. We investigated the interaction between body movements and the flow around swimming eels using two-dimensional particle image velocimetry. Maximum flow velocities adjacent to the eel9s body increase almost linearly from head to tail, suggesting that eels generate thrust continuously along their body. The wake behind eels swimming at 1.5Ls-1, where L is body length, consisted of a double row of double vortices with little backward momentum. The eel sheds two vortices per half tail-beat, which can be identified by their shedding dynamics as a start—stop vortex of the tail and a vortex shed when the body-generated flows reach the `trailing edge9 and cause separation. Two consecutively shed ipsilateral body and tail vortices combine to form a vortex pair that moves away from the mean path of motion. This wake shape resembles flow patterns described previously for a propulsive mode in which neither swimming efficiency nor thrust is maximised but sideways forces are high. This swimming mode is suited to high manoeuvrability. Earlier recordings show that eels also generate a wake reflective of maximum swimming efficiency. The combined findings suggest that eels can modify their body wave to generate wakes that reflect their propulsive mode.

Suppression of aggression in rainbow trout (Oncorhynchus mykiss) by dietary l-tryptophan

Abstract: SUMMARY Juvenile rainbow trout Oncorhynchus mykiss were isolated in individual compartments in observation aquaria and allowed to acclimate for 1 week, during which they were fed commercial trout feed. Thereafter, the fish were tested for aggressive behaviour using a resident/intruder test. Following this first resident/intruder test, the feed was exchanged for an experimental wet feed supplemented with 0.15 % or 1.5 % l-tryptophan (by wet mass). Controls received the same feed but without l-tryptophan supplementation. The fish were fed to satiety daily, and their individual feed intake was recorded. Aggressive behaviour was quantified again after 3 and 7 days of l-tryptophan feeding using the resident/intruder test. Feeding the fish l-tryptophan-supplemented feed for 3 days had no effect on aggressive behaviour, whereas feeding the fish l-tryptophan-supplemented feed for 7 days significantly suppressed aggressive behaviour in the fish, an effect seen at both levels of l-tryptophan supplementation. Fish fed l-tryptophan-supplemented feed showed elevated plasma and brain levels of l-tryptophan. The amino acid l-tryptophan is the precursor of serotonin, and supplementary dietary l-tryptophan was found to elevate levels of 5-hydroxyindoleacetic acid (5-HIAA) and the 5-HIAA/serotonin concentration ratio in the brain. Neither feed intake nor plasma cortisol level was significantly affected by dietary l-tryptophan. Central serotonin is believed to have an inhibitory effect on aggressive behaviour, and it is suggested that the suppressive effect of dietary l-tryptophan on aggressive behaviour is mediated by an elevation of brain serotonergic activity.

Hindlimb muscle function in relation to speed and gait: in vivo patterns of strain and activation in a hip and knee extensor of the rat (Rattus norvegicus)

Abstract: Understanding how animals actually use their muscles during locomotion is an important goal in the fields of locomotor physiology and biomechanics. Active muscles in vivo can shorten, lengthen or remain isometric, and their mechanical performance depends on the relative magnitude and timing of these patterns of fascicle strain and activation. It has recently been suggested that terrestrial animals may conserve metabolic energy during locomotion by minimizing limb extensor muscle strain during stance, when the muscle is active, facilitating more economical force generation and elastic energy recovery from limb muscle–tendon units. However, whereas the ankle extensors of running turkeys and hopping wallabies have been shown to generate force with little length change (<6% strain), similar muscles in cats appear to change length more substantially while active. Because previous work has tended to focus on the mechanical behavior of ankle extensors during animal movements, the actions of more proximal limb muscles are less well understood. To explore further the hypothesis of force economy and isometric behavior of limb muscles during terrestrial locomotion, we measured patterns of electromyographic (EMG) activity and fascicle strain (using sonomicrometry) in two of the largest muscles of the rat hindlimb, the biceps femoris (a hip extensor) and vastus lateralis (a knee extensor) during walking, trotting and galloping. Our results show that the biceps and vastus exhibit largely overlapping bursts of electrical activity during the stance phase of each step cycle in all gaits. During walking and trotting, this activity typically commences shortly before the hindlimb touches the ground, but during galloping the onset of activity depends on whether the limb is trailing (first limb down) or leading (second limb down), particularly in the vastus. In the trailing limb, the timing of the onset of vastus activity is slightly earlier than that observed during walking and trotting, but in the leading limb, this activity begins much later, well after the foot makes ground contact (mean 7% of the step cycle). In both muscles, EMG activity typically ceases approximately two-thirds of the way through the stance phase. While electrically active during stance, biceps fascicles shorten, although the extent of shortening differs significantly among gaits ( P <0.01). Total average fascicle shortening strain in the biceps is greater during walking (23±3%) and trotting (27±5%) than during galloping (12±5% and 19±6% in the trailing and leading limbs, respectively). In contrast, vastus fascicles typically lengthen (by 8–16%, depending on gait) over the first half of stance, when the muscle is electrically active, before shortening slightly or remaining nearly isometric over much of the second half of stance. Interestingly, in the leading limb during galloping, vastus fascicles lengthen prior to muscle activation and exhibit substantial shortening (10±2%) during the period when EMG activity is recorded. Thus, patterns of muscle activation and/or muscle strain differ among gaits, between muscles and even within the same muscle of contralateral hindlimbs (as during galloping). In contrast to the minimal strain predicted by the force economy hypothesis, our results suggest that proximal limb muscles in rats operate over substantial length ranges during stance over various speeds and gaits and exhibit complex and changing activation and strain regimes, exemplifying the variable mechanical roles that muscles can play, even during level, steady-speed locomotion.

Evolution of water balance in the genus Drosophila

Abstract: SUMMARY Fruit flies of the genus Drosophila have independently invaded deserts around the world on numerous occasions. To understand the physiological mechanisms allowing these small organisms to survive and thrive in arid environments, we performed a phylogenetic analysis of water balance in Drosophila species from different habitats. Desert (cactophilic) species were more resistant to desiccation than mesic ones. This resistance could be accomplished in three ways: by increasing the amount of water in the body, by reducing rates of water loss or by tolerating the loss of a greater percentage of body water (dehydration tolerance). Cactophilic Drosophila lost water less rapidly and appeared to be more tolerant of low water content, although males actually contained less water than their mesic congeners. However, when the phylogenetic relationships between the species were taken into account, greater dehydration tolerance was not correlated with increased desiccation resistance. Therefore, only one of the three expected adaptive mechanisms, lower rates of water loss, has actually evolved in desert Drosophila , and the other apparently adaptive difference between arid and mesic species (increased dehydration tolerance) instead reflects phylogenetic history.

Dependence of human squat jump performance on the series elastic compliance of the triceps surae: a simulation study.

Abstract: The purposes of this study were to determine the dependence of human squat jump performance on the compliance of series elastic elements (SEEs) of the triceps surae (consisting of the soleus and gastrocnemius) and to explain this dependence. Vertical squat jumps were simulated using an optimal control model of the human musculo-skeletal system. Maximum jump height was found for several values of triceps surae SEE strain at maximum isometric force (e (0)). When e (0) was increased from 1 to 10 %, maximum jump height increased by 8 cm. This was partly due to a higher work output of contractile elements (CEs) of the muscles, primarily of the soleus, and also partly to an increased efficacy of converting muscle work to energy contributing to jump height. The soleus produced more work at e (0)=10 % because, as a result of SEE recoil, the CE covered its shortening range at lower velocity and hence produced more force. Efficacy was higher at e (0)=10 % because a higher vertical velocity at take-off was achieved with a lower rotational energy of the body segments. This apparent discrepancy was explained by increased angular velocities of the shanks and feet, which have small moments of inertia, and decreased angular velocities of the thighs and trunk, which have larger moments of inertia. This redistribution of segmental contributions to the vertical velocity of the centre of mass was possible because the increased compliance of the triceps surae SEE enhanced the energy-buffering capacity of this muscle group and, thereby, allowed for a higher power output at the ankles. It seems that long compliant tendons in the plantar flexors are an elegant solution to the problem of maximizing jumping performance.

Limits to sustained energy intake: I. Lactation in the laboratory mouse MUS MUSCULUS

Abstract: SUMMARY Laboratory mice (strain MF1) were used to determine whether sustainable rates of energy intake are limited during lactation. Mice raising natural-sized litters ( N =71) reached an asymptote in their daily food intake between days 13 and 16 of lactation at 23.1gday −1 and also between litter sizes of 9 and 15 pups (22.8gday −1 ). A second group of 37 females had their litter sizes manipulated at birth to raise more or fewer offspring than they gave birth to. When the litter size was increased, females did not increase their food intake to match their new litter size. However, when litter size was decreased, females decreased their asymptotic daily food intake during late lactation in relation to the extent of reduction in litter size. Therefore, it appeared that females were limited during late lactation and with large litter sizes. The milk energy exported amounted to 44% of the gross energy intake, and the estimated daily energy expenditure was therefore considerably lower than the sustained energy intake [8.0×RMR(gross), 6.6×RMR(assimilated)], and averaged 3.1×RMR, where RMR is resting metabolic rate. It was not possible to determine whether the apparent limit on sustained energy intake was acting centrally or peripherally because of the asymptotes in both food intake and milk energy output with increasing litter size.

Evaluating the use of ram and suction during prey capture by cichlid fishes

Abstract: We characterized prey-capture strategies in seven species of cichlid fishes representing diverse trophic habits and anticipated feeding abilities. The species examined were Petenia splendida , Cichla ocellaris , Cichlasoma minckleyi , Astronotus ocellatus , Crenicichla geayi , Heros severus (formerly Cichlasoma severum ) and Cyprichromis leptosoma . Three individuals per species were filmed with video at 500Hz as they captured live adult Artemia sp. and Poecilia reticulata . For each feeding sequence, we measured the contribution of predator movement towards the prey (i.e. ram) and the movement of prey towards the predator due to suction. The use of ram differed significantly among prey types and predator species, varying as much as sixfold across predator species. High values of ram resulted in high attack velocities. Jaw protrusion contributed as much as 50% to overall ram values in some species, verifying its role in enhancing attack velocity. Suction distance did not vary significantly among species. Diversity in prey-capture behavior was therefore found to reflect differences among species in the strategy used to approach prey. Limited variation in the distance from which prey were sucked into the mouth is interpreted as the result of an expected exponential decline in water velocity with distance from the mouth of the suction-feeding predator. We propose that this relationship represents a major constraint on the distance over which suction feeding is effective for all aquatic-feeding predators.

Prey-capture behavior in gymnotid electric fish: motion analysis and effects of water conductivity.

Abstract: Animals can actively influence the content and quality of sensory information they acquire from the environment through the positioning of peripheral sensory surfaces. This study investigated receptor surface positioning during prey-capture behavior in weakly electric gymnotiform fish of the genus Apteronotus. Infrared video techniques and three-dimensional model-based tracking methods were used to provide quantitative information on body position and conformation as black ghost (A. albifrons) and brown ghost (A. leptorhynchus) knifefish hunted for prey (Daphnia magna) in the dark. We found that detection distance depends on the electrical conductivity of the surrounding water. Best performance was observed at low water conductivity (2.8 cm mean detection distance and 2 % miss rate at 35 microS cm(−)(1), A. albifrons) and poorest performance at high conductivity (1.5 cm mean detection distance and 11 % miss rate at 600 microS cm(−)(1), A. albifrons). The observed conductivity-dependence implies that nonvisual prey detection in Apteronotus is likely to be dominated by the electrosense over the range of water conductivities experienced by the animal in its natural environment. This result provides the first evidence for the involvement of electrosensory cues in the prey-capture behavior of gymnotids, but it leaves open the possibility that both the high-frequency (tuberous) and low-frequency (ampullary) electroreceptors may contribute. We describe an electrosensory orienting response to prey, whereby the fish rolls its body following detection to bring the prey above the dorsum. This orienting response and the spatial distribution of prey at the time of detection highlight the importance of the dorsal surface of the trunk for electrosensory signal acquisition. Finally, quantitative analysis of fish motion demonstrates that Apteronotus can adapt its trajectory to account for post-detection motion of the prey, suggesting that it uses a closed-loop adaptive tracking strategy, rather than an open-loop ballistic strike strategy, to intercept the prey.

Effects of high intensity exercise training on cardiovascular function, oxygen uptake, internal oxygen transport and osmotic balance in chinook salmon (Oncorhynchus tshawytscha) during critical speed swimming.

Abstract: SUMMARY To examine cardiorespiratory plasticity, cardiovascular function, oxygen consumption, oxygen delivery and osmotic balance were measured at velocities up to critical swimming speed ( U crit ) in seawater-adapted chinook salmon. We used two groups of fish. The control group had swum continuously for 4 months at a low intensity (0.5 BL s -1 ) and the other was given a high-intensity training regimen (a U crit swim test on alternate days) over the same period of time. Compared with available data for other salmonid species, the control group had a higher maximum oxygen consumption ( Ṁ o 2max ; 244μ mol O 2 min -1 kg -1 ), cardiac output ( Q max ; 65 ml min -1 kg -1 ) and blood oxygen content ( C a O 2 ; 15 ml O 2 dl -1 ). Exercise training caused a 50% increase in Ṁ o 2max without changing either U crit or C a O 2 , even though there were small but significant increases in hematocrit, hemoglobin concentration and relative ventricular mass. During swimming, however, exercise-trained fish experienced a smaller decrease in body mass and muscle moisture, a smaller increase in plasma osmolality, and reduced venous oxygen stores compared with control fish. Consequently, exercise training apparently diminished the osmo—respiratory compromise, but improved oxygen extraction at the tissues. We conclude that the training-induced increase in Ṁ o 2max provided benefits to systems other than the locomotory system, such as osmoregulation, enabling trained fish to better multitask physiological functions while swimming. Furthermore, because a good interspecific correlation exists between Ṁ o 2max and arterial oxygen supply ( Ṫ o 2max ; r 2 =0.99) among temperate fish species, it is likely that C a O 2 and Q max are principal loci for cardiorespiratory evolutionary adaptation but not for intraspecific cardiorepiratory plasticity as revealed by high intensity exercise training.

Swing- and support-related muscle actions differentially trigger human walk–run and run–walk transitions

Abstract: SUMMARY There has been no consistent explanation as to why humans prefer changing their gait from walking to running and from running to walking at increasing and decreasing speeds, respectively. This study examined muscle activation as a possible determinant of these gait transitions. Seven subjects walked and ran on a motor-driven treadmill for 40s at speeds of 55, 70, 85, 100, 115, 130 and 145% of the preferred transition speed. The movements of subjects were videotaped, and surface electromyographic activity was recorded from seven major leg muscles. Resultant moments at the leg joints during the swing phase were calculated. During the swing phase of locomotion at preferred running speeds (115, 130, 145%), swing-related activation of the ankle, knee and hip flexors and peaks of flexion moments were typically lower ( P P