Social structure evolves from a trade-off between the costs and benefits of group-living, which are in turn dependent upon the distribution of key resources such as food and shelter. Males and females, or juveniles and adults, may have different priorities when selecting habitat due to differences in physiological or behavioural imperatives, leading to complex patterns in group composition. We studied social structure and mating behaviour in the insectivorous bat Myotis daubentonii along an altitudinal gradient, combining field studies with molecular genetics. With increasing altitude the proportion of males in summer roosts increased and only males were present in the highest roosts. With increasing altitude environmental temperature decreased, nightly variation in temperature increased, and bat foraging activity decreased, supporting the hypothesis that the harsher, high elevation sites cannot support breeding females. We found that offspring in female-dominated lowland roosts had a very high probability of being fathered by bats caught during autumnswarming at hibernation sites, in contrast to those in intermediate roosts, which had a high probability of being fathered by males sharing the nursery roost with the females. Whilst females normally appear to exclude males from nursery colonies, for those in marginal habitats, one explanation for the presence of males is that the thermoregulatory benefits to the females may outweigh disadvantages, such as competition for food, and give some males an opportunity to increase their breeding success. We suggest that the environment, and its effects on resource distribution, thus determine social structure, which in turn determines the mating pattern that has evolved.
hide

Including predictors in species distribution models at inappropriate spatial scales can decrease the variance explained, add residual spatial autocorrelation (RSA) and lead to the wrong conclusions. Some studies have measured predictors within different buffer sizes (scales) around sample locations, regressed each predictor against the response at each scale and selected the scale with the best model fit as the appropriate scale for this predictor. However, a predictor can influence a species at several scales or show several scales with good model fit due to a bias caused by RSA. This makes the evaluation of all scales with good model fit necessary. With potentially several scales per predictor and multiple predictors to evaluate, the number of predictors can be large relative to the number of data points, potentially impeding variable selection with traditional statistical techniques, such as logistic regression. We trialled a variable selection process using the random forest algorithm, which allows the simultaneous evaluation of several scales of multiple predictors. Using simulated responses, we compared the performance of models resulting from this approach with models using the known predictors at arbitrary and at the known spatial scales. We also apply the proposed approach to a real data set of curlew (Numenius arquata). AIC, AUC and Naglekerke's pseudo R of the models resulting from the proposed variable selection were often very similar to the models with the known predictors at known spatial scales. Only two of nine models required the addition of spatial eigenvectors to account for RSA. Arbitrary scale models always required the addition of spatial eigenvectors. 75% (50-100%) of the known predictors were selected at scales similar to the known scale (within 3 km). In the curlew model, predictors at large, medium and small spatial scales were selected, suggesting that for appropriate landscape-scale models multiple scales need to be evaluated. The proposed approach selected several of the correct predictors at appropriate spatial scales out of 544 possible predictors. Thus, it facilitates the evaluation of multiple spatial scales of multiple predictors against each other in landscape-scale models.© 2012 The Authors. Methods in Ecology and Evolution © 2012 British Ecological Society.
hide

A successful and safe methodology for the subcutaneous insertion of passive integrated transponder (PIT) tags in a small- to medium-sized bat (average mass 9 g) under isoflurane-induced anaesthesia is described. Passive integrated transponder (PIT) tagging had no significant impact on the rate of recapture, body condition index (BCI) (bodyweight/forearm length) and reproductive success of tagged individuals, and no visible injuries or health problems were observed in any of the recaptured bats. Tagging success, in terms of retention and function, was 92 per cent (n=61) by the third year of using the method. Sixteen per cent (n=39) of bats tagged during the three-year study period were not producing positive scans with the microchip reader when recaptured after previously successful tag insertion, indicating that the tags were either working their way out of the bats or ceasing to function.
hide

Comparative ecological and behavioural studies of the widespread and diverse Vespertilionidae, which comprise almost 400 of the 1100 bat species, have been limited by the availability of markers. The potential of new methods for developing conserved microsatellite markers that possess enhanced cross-species utility has recently been illustrated in studies of birds. We have applied these methods to develop enhanced microsatellite markers for vespertilionid bats, in particular for the genus Myotis (103 species). We compared published bat microsatellites with their homologues in the genome sequence of the little brown bat, Myotis lucifugus, to create consensus sequences that were used to design candidate primer sets. Primer sets were then tested for amplification and polymorphism in 22 species of bat from nine of the largest families (including 11 Vespertilionidae). Of 46 loci tested, 33 were polymorphic, on average, in each of seven Myotis species tested, 20 in each of four species in other vespertilionid genera, and two in 11 nonvespertilionid species.© 2012 Blackwell Publishing Ltd.
hide

Major roads can reduce bat abundance and diversity over considerable distances. To mitigate against these effects and comply with environmental law, many European countries install bridges, gantries or underpasses to make roads permeable and safer to cross. However, through lack of appropriate monitoring, there is little evidence to support their effectiveness. Three underpasses and four bat gantries were investigated in northern England. Echolocation call recordings and observations were used to determine the number of bats using underpasses in preference to crossing the road above, and the height at which bats crossed. At gantries, proximity to the gantry and height of crossing bats were measured. Data were compared to those from adjacent, severed commuting routes that had no crossing structure. At one underpass 96% of bats flew through it in preference to crossing the road. This underpass was located on a pre-construction commuting route that allowed bats to pass without changing flight height or direction. At two underpasses attempts to divert bats from their original commuting routes were unsuccessful and bats crossed the road at the height of passing vehicles. Underpasses have the potential to allow bats to cross roads safely if built on pre-construction commuting routes. Bat gantries were ineffective and used by a very small proportion of bats, even up to nine years after construction. Most bats near gantries crossed roads along severed, pre-construction commuting routes at heights that put them in the path of vehicles. Crossing height was strongly correlated with verge height, suggesting that elevated verges may have some value in mitigation, but increased flight height may be at the cost of reduced permeability. Green bridges should be explored as an alternative form of mitigation. Robust monitoring is essential to assess objectively the case for mitigation and to ensure effective mitigation.
hide

Plecotus auritus, a small, gleaning bat species, lives in small, isolated summer colonies in which both males and females show a high degree of natal philopatry. Despite this, colonies have high gene diversities and low inbreeding coefficients. It has been suggested that inbreeding is avoided because mating occurs during autumnal and spring swarming at hibernation sites. We tested this hypothesis by comparing microsatellite profiles, based on eight loci, of bats from six summer colonies and two swarming sites they were known to visit from radiotelemetry studies. We found high gene diversities (H (s) = 0.77) at both swarming sites and summer colonies which were not statistically different. There was no detectable isolation by distance and F-ST was low (0.001). Together, these results suggest high gene flow between sites. Despite this, there was small but significant genetic differentiation amongst summer colonies and between summer colonies and the primary swarming site. We suggest that swarming is important for gene flow and for maintaining genetic diversity in this highly philopatric species and discuss possible reasons for the genetic differentiation observed. The identification and protection of swarming sites should be a major conservation priority for this and other temperate bat species.
hide

Eels are capable of locomotion both in water and on land using undulations of the body axis. Axial undulations are powered by the lateral musculature. Differences in kinematics and the underlying patterns of fast muscle activation are apparent between locomotion in these two environments. The change in isometric fast muscle properties with axial location was less marked than in most other species. Time from stimulus to peak force (T(a)) did not change significantly with axial position and was 82+/-6 ms at 0.45BL and 93+/-3 ms at 0.75BL, where BL is total body length. Time from stimulus to 90% relaxation (T(90)) changed significantly with axial location, increasing from 203+/-11ms at 0.45BL to 239+/-9 ms at 0.75BL. Fast muscle power outputs were measured using the workloop technique. Maximum power outputs at +/-5% strain using optimal stimuli were 17.3+/-1.3W kg(-1) in muscle from 0.45BL and 16.3+/-1.5W kg(-1) in muscle from 0.75BL. Power output peaked at a cycle frequency of 2Hz. The stimulus patterns associated with swimming generated greater force and power than those associated with terrestrial crawling. This decrease in muscle performance in eels may occur because on land the eel is constrained to a particular kinematic pattern in order to produce thrust against an underlying substratum.
hide

Strain and activity patterns were determined during slow steady swimming (tailbeat frequency 1.5-2.5 Hz) at three locations on the body in the slow myotomal muscle of rainbow trout Oncorhynchus mykiss using sonomicrometry and electromyography. Strain was independent of tailbeat frequency over the range studied and increased significantly from +/-3.3% l(0) at 0.35BL to +/-6% at 0.65BL, where lo is muscle resting length and BL is total body length. Muscle activation occurred significantly later in the strain cycle at 0.35BL (phase shift 59 degrees) than at 0.65BL (30 degrees), and the duration of activity was significantly longer (211 degrees at 0.35BL and 181 degrees at 0.65BL), These results differ from those of previous studies. The results have been used to simulate in vivo activity in isolated muscle preparations using the work loop technique. Preparations from all three locations generated net positive power under in vivo conditions, but the negative power component increased from head to tail. Both kinematically, and in the way its muscle functions to generate hydrodynamic thrust, the rainbow trout appears to be intermediate between angulliform swimmers such as the eel, which generate thrust along their entire body length, and carangiform fish (e.g. saithe Pollachius virens), which generate thrust primarily at the tail blade.
hide

It has been hypothesised that regional endothermy has evolved in the muscle of some tunas to enhance the locomotory performance of the fish by increasing muscle power output, Using the work loop technique, we have determined the relationship between cycle frequency and power output, over a range of temperatures, in isolated bundles of slow muscle fibres from the endothermic yellowfin tuna (Thunnus albacares) and its ectothermic relative the bonito (Sai da chiliensis), Power output in all preparations was highly temperature-dependent. A counter-current heat exchanger which could maintain a 10 degrees C temperature differential would typically double maximum muscle power output and the frequency at which maximum power is generated (f(opt)) The deep slow muscle of the tuna was able to operate at higher temperatures than slow muscle from the bonito, but was more sensitive to temperature change than more superficially located slow fibres from both tuna and bonito, This suggests that it has undergone some evolutionary specialisation for operation at higher, but relatively stable, temperatures, f(opt) of slow muscle was higher than the tailbeat frequency of undisturbed cruising tuna and, together with the high intrinsic power output of the slow muscle mass, suggests that cruising fish have a substantial slow muscle power reserve, This reserve should be sufficient to power significantly higher sustainable swimming speeds presumably at lower energetic cost than if intrinsically less efficient fast fibres were recruited.
hide

The work loop technique was used to measure the mechanical performance in situ of the latissimus dorsi (LD) muscles of rabbits maintained under fentanyl anesthesia. After 3 wk of incrementally applied stretch the LD muscles were 36% heavier, but absolute power output (195 mW/muscle) was not significantly changed relative to that of external control muscle (206 mW). In contrast, continuous 10-Hz electrical stimulation reduced power output per kilogram of muscle>75% after 3 or 6 wk and muscle mass by 32% after 6 wk. When combined, stretch and 10-Hz electrical stimulation preserved or increased the mass of the treated muscles but failed to prevent an 80% loss in maximum muscle power. However, this combined treatment increased fatigue resistance to a greater degree than electrical stimulation alone. These stretched/stimulated muscles, therefore, are more suitable for cardiomyoplasty. Nonetheless, further work will be necessary to find an ideal training program for this surgical procedure.
hide

The power output of rabbit latissimus dorsi muscle was calculated under isotonic conditions and during oscillatory work. Isotonic shortening studies yielded a maximum power output of 120 W . kg-1 at a P/P0 of 0.4 compared to a maximum power output of 32 W . kg-1 obtained using the work loop technique. This difference can largely be explained by comparing actual work loops with those constructed using force velocity (P/V) and isometric data. At low cycle frequencies, work loop power output is quite close to that predicted from P/V and isometric data. However, at higher frequencies other dynamic muscle properties appear to exert a more marked effect.
hide

The European pipistrelle, Pipistrellus pipistrellus, consists of two phonic types that echolocate with frequencies of maximum energy (FMAXE) averaging 46 kHz and 55 kHz (named the 45 kHz and 55 kHz phonic types for simplicity). These two phonic types occur in sympatry over much of Britain. Between 1993 and 1995 mating groups of P. pipistrellus were located in bat boxes between August and October, the main mating period for these bats. Mating groups comprised a single male and up to three females. Broadband, time-expanded recordings of search-phase echolocation calls were made from bats immediately after release. The average FMAXE of each bat was calculated: bats with an FMAXE less than 49 kHz were classed as 45 kHz phonic types and bats with an FMAXE greater than 52 kHz were classed as 55 kHz phonic types. A total of 16, 45 kHz groups and ten 55 kHz groups were located. No mating group ever contained more than one phonic type, although both types were known to be present at several of the study sites. Males emitted calls of higher frequency than females for both phonic types. The study suggests that the two phonic types are reproductively isolated and supports the hypothesis that P. pipistrellus should be considered as two cryptic sibling species.
hide

Trichinella spiralis larvae infect and develop within skeletal muscle cells causing major changes to their mechanical properties. The aim of this investigation was to determine the effects of T. spiralis on the power output and fatigue resistance of the mammalian diaphragm under conditions simulating in vivo operation and to relate these to respiratory performance. Infection with T. spiralis leads to major reductions in mechanical stress, work, power output and fatigue resistance. These changes are associated with the number of larvae present in the muscle and the duration of infection. However, the initial decline in mechanical performance occurs during the onset of infection when there are few larvae observed within the muscle cells, indicating that T. spiralis may affect the properties of muscle before encapsulation. This may correspond to the host's inflammatory response and the effects of larval excretory/secretory products. The decline in mechanical performance will have a profound effect on respiration both at rest and during exertion. This must influence the behaviour of the host and increase its chance of capture by predators, which is likely to benefit the parasite by facilitating its transmission.
hide

The mechanical properties of soleus and extensor digitorum longus (EDL) muscles from the mouse were studied using the work loop technique. Under optimum conditions, the EDL produced a maximum mean power output of 107 W kg-1 at a cycle frequency of 10 Hz. In comparison, the maximum mean power output of the soleus was 34 W kg-1 at 5 Hz cycle frequency. Video analysis of mice determined the stride frequency range to be from 2.87 Hz at a walk to 8.23 Hz at a flat-out gallop, with the trot-to-gallop transition occurring at 5.89 Hz. In vivo EDL electromyogram (EMG) activity is recorded primarily during shortening and the muscle operates in a power-generating mode. The soleus is close to isometric when EMG activity is recorded, but mechanical activity persists into the shortening phase. Both muscles are likely to operate over cycle frequency ranges just below, or at, those yielding maximal power. Soleus and EDL produced maximal power output in vitro when operating at mean sarcomere lengths of 2.58 microns and 2.71 microns respectively. These lengths are slightly above the plateau of the length-force curve predicted for rat leg muscle (2.3-2.5 microns). The sarcomere length ranges used in vivo by the soleus and EDL were determined, by fixing muscles in the extreme active positions predicted from video and cine analysis, to be 2.28-2.57 microns and 2.49-2.88 microns respectively. These ranges are both close to those shown to yield maximum power output in vitro and to the plateau of the sarcomere length-force curve.
hide

The influence of length on work production was investigated for rat papillary muscles using the work loop technique. Active and passive length-force relationships were first determined under isometric conditions and the length for maximum force production (Lmax) was derived. Starting from different lengths within the physiological range, a series of work loops was generated using the stimulation phase shift, strain amplitude and cycle frequency previously found to be optimal for power output at 37 degrees C. The relationship between muscle length and net work was used to determine the length at which work output was maximal (Lopt). In order to examine the dynamic passive properties of the muscles, unstimulated muscles were subjected to the same regime of sinusoidal oscillation as used for the active loops. From the hysteresis loops, lengthening work (work done to extend the passive muscle), passive shortening work (work returned during shortening) and net energy loss (hysteresis) could be measured. The decline in net work production at lengths greater than 95% Lmax could largely be attributed to the rapid and non-linear increase in muscle stiffness and the increase in net energy loss over this range of lengths. The physiological significance of the length-work relationship is considered and the mechanical properties of active and passive papillary muscles are discussed with reference to sarcomere length and cardiac muscle ultrastructure.
hide

Papillary muscles were isolated from the right ventricles of rats and the length for maximum active force generation (Lmax) was determined isometrically. The work loop technique was used to derive the length for maximum work production (Lopt) at the cycle frequency, strain amplitude and stimulation phase shift found to be optimal for power output. Lopt was typically 7% shorter than Lmax and within the physiological length range (87.5% Lmax to Lmax). Net work and power output were measured during sinusoidal strain cycles around Lopt, over the cycle frequency range 1-9 Hz, strain amplitude and phase shift being optimised for work and power at each frequency. Experiments were performed at 37 degrees C. Distinct optima were found in both the work-frequency and the power-frequency relationships. The optimum cycle frequency for net work production was lower than the frequency for maximum power output. The mean maximum power output at 37 degrees C was 8.62 +/- 0.50 W kg-1 (mean +/- S.E.M., N = 9) and was achieved at a cycle frequency of approximately 6 Hz, close to the estimated resting heart rate of 5.8 Hz for the rats used (mean mass 223 +/- 25 g). The cycle frequency, strain amplitude and stimulation phase shift found to be optimal for power output produced an in vitro contraction closely simulating the basal in vivo contraction.
hide

Papillary muscles were isolated from the right ventricles of rats and the length for maximum active force generation (Lmax) was determined isometrically. The work loop technique was used to derive the length for maximum work production (Lopt) at the cycle frequency, strain amplitude and stimulation phase shift found to be optimal for power output. Lopt was typically 7% shorter than Lmax and within the physiological length range (87.5% Lmax to Lmax). Net work and power output were measured during sinusoidal strain cycles around Lopt, over the cycle frequency range 1-9 Hz, strain amplitude and phase shift being optimised for work and power at each frequency. Experiments were performed at 37 degrees C. Distinct optima were found in both the work-frequency and the power-frequency relationships. The optimum cycle frequency for net work production was lower than the frequency for maximum power output. The mean maximum power output at 37 degrees C was 8.62 +/- 0.50 W kg-1 (mean +/- S.E.M., N = 9) and was achieved at a cycle frequency of approximately 6 Hz, close to the estimated resting heart rate of 5.8 Hz for the rats used (mean mass 223 +/- 25 g). The cycle frequency, strain amplitude and stimulation phase shift found to be optimal for power output produced an in vitro contraction closely simulating the basal in vivo contraction.
hide

The isometric mechanical properties of diaphragm muscle were studied in mice infected with either Trichinella spiralis or Trichinella pseudospiralis. Measurements of muscle stress were taken at 15, 20, 30 and 40 days post-infection. Infected diaphragm muscle showed a significant (P<0.001) reduction in muscle stress during both twitch and tetanic contractions when compared with muscle from control animals. T. spiralis caused a significant reduction in resistance to muscle fatigue. The consequences of these changes in muscle function to host pathology are discussed, and related to previous work on the effects of Trichinella on host biochemistry and the immune response.
hide

Video and ciné films of mammals running at the trot-gallop transition were analysed to measure breathing frequencies. Breathing frequency at the trot-gallop transition (fb, in Hz) was shown to decrease with increasing body mass (M, in kg) and was described by the equation fb = 5.08 M-0.14. The stiffness of thethorax and diaphragm of mice, rats, rabbits and wallabies was calculated and this, together with the mass of the viscera, was used to calculate the natural frequency of the system (nft, in Hz). The relationship between nft and body mass can be described by the equation nft = 5.02M-0.18. The significance of these results is discussed in relation to models of mechanical linkage between respiratory and locomotory movements.
hide

Bundles of muscle fibres were isolated from the diaphragm of mouse, rat and rabbit. Mean oscillatory power output was determined during phasic stimulation and imposed sinusoidal length changes. Maximum power output was measured over a range of cycle frequencies. The cycle frequency for maximum power output (fopt) decreased with increasing body mass and was described by the equation, fopt = 4.42M-0.16, where M is body mass. A very similar relationship has been reported between body mass and the frequency of the trot-gallop transition in terrestrial, quadrupedal mammals [Heglund et al. (1974), Science 186, 1112-1113), and the significance of this similarity is discussed.
hide

Fast myotomal muscle fibres were isolated from the cod (Gadus morhua L.) and the energy cost of contraction was measured under conditions simulating swimming. Fibre bundles were subjected to sinusoidal cycles of shortening and lengthening about their in situ fibre length, and stimulated at selected phases in each cycle. The preparations were poisoned with iodoacetic acid and bubbled with nitrogen to block the synthesis of ATP. After an initial rapid decline over the first 10 cycles, force and net work remained steady in some cases for up to 64 oscillatory length cycles, but more commonly declined slowly after about 30 cycles. The total mechanical work performed increased largely in proportion to the number of work cycles. At the end of each experiment fibres were frozen in isopentane cooled in liquid nitrogen and metabolite concentrations determined by high performance liquid chromatography (HPLC) and enzymatic analysis. Concentrations of adenylates did not differ significantly from control values, although a significant increase in IMP concentrations at 64 cycles accounted for the maintenance of relatively high energy charge values. Creatine (C) concentrations increased and creatine phosphate (CP) concentrations decreased, implying a tight coupling of the ATP/ADP reaction to the CP/C reaction. Muscle economy was calculated as the positive work performed during a work cycle divided by the total chemical energy expended. These values (approx. 7 mJ mumol-1) were found to be independent of the number of work cycles performed, although a trend to increase was observed. Muscle efficiency values, calculated assuming a Gibb's force free energy change for CP splitting in vivo of 55 kJ mol-1, were in the range 12-23%.
hide

Single or small bundles of fibres were isolated from the abdominal myotomes of the sculpin Myoxocephalus scorpius, a teleost with a polyneuronal pattern of fast muscle innervation. Fibres responded to a supra-threshold stimulus with an all-or-none twitch. Tetanic fusion frequency at 3 degrees C was 40-60 Hz, and the twitch tetanus ratio 0.70. Maximum isometric tension was 281 kN m-2. Similar isometric contractile properties were obtained from the focally innervated fast muscle fibres of another teleost, the eel, Anguilla anguilla. The response of sculpin fibres to stretch during tetanus was similar to that reported for frog twitch fibres. A 5% stretch of 25-50 ms duration increased force to 1.4 Po which decayed to a steady level 5-10% above that of a control tetanus. The force-velocity relationship was also studied. Maximum contraction velocity was 4.75 Ls-1. Force-velocity data were not adequately described by a simple hyperbola. Alternative methods of curve fitting have been explored and discussed.
hide