Costs of encephalization: the energy trade-off hypothesis tested on birds

Costs and benefits of encephalization are a major topic of debate in the study of primate and human evolution. Comparative studies provide an opportunity to test the validity of a hypothesis as a general principle, rather than it being a special case in primate or hominid evolution. If a population evolves a larger brain, the metabolic costs of doing so must be paid for by either an increased energy turnover (direct metabolic constraint) or by a trade-off with other energetically expensive costs of body maintenance, locomotion, or reproduction, here referred to as the energy trade-off hypothesis, an extension of the influential Expensive Tissue Hypothesis of Aiello and Wheeler (1995, Curr. Anthropol. 36, 199-221). In the present paper, we tested these hypotheses on birds using raw species values, family means, and independent contrasts analysis to account for phylogenetic influences. First, we tested whether basal metabolic rates are correlated with brain mass or any other variable of interest. This not being the case, we examined various trade-offs between brain mass and the mass of other expensive tissues such as gut mass, which is approximated by gut length or diet quality. Only weak support was found for this original Expensive Tissue Hypothesis in birds. However, other energy allocations such as locomotor mode and reproductive strategy may also be reduced to shunt energy to an enlarged brain. We found a significantly negative correlation between brain mass and pectoral muscle mass, which averages 18% of body mass in birds and is indicative of their relative costs of flight. Reproductive costs, on the other hand, are positively correlated with brain mass in birds. An increase in brain mass may allow birds to devote more energy to reproduction, although not through an increase in their own energy budget as in mammals, but through direct provisioning of their offspring. The trade-off between locomotor costs and brain mass in birds lets us conclude that an analogous effect could have played a role in the evolution of a larger brain in human evolution.     

Directed flight and optimal airspeeds: homeward‐bound gulls react flexibly to wind yet fly slower than predicted

Birds in flight are proposed to adjust their body orientation (heading) and airspeed to wind conditions adaptively according to time and energy constraints. Airspeeds in goal‐directed flight are predicted to approach or exceed maximum‐range airspeeds, which minimize transport costs (energy expenditure per unit distance) and should increase in headwinds and crosswinds. Diagnosis of airspeed adjustment is however obscured by uncertainty regarding birds' goal‐directions, transport costs, interrelations with orientation strategy and the attainability of predicted behaviour. To address these issues, we tested whether gulls minimized transport costs through adjustment of airspeed and heading to wind conditions during extended inbound flight over water (180–360 km) to their breeding colony, and introduce a methodology to assess transport (energy) efficiency given wind conditions. Airspeeds, heading, flight mode and energy expenditure were estimated using GPS tracking, accelerometer and wind data. Predicted flight was determined by simulating each trip according to maximum‐range airspeeds and various orientation strategies. Gulls employed primarily flapping flight (93%), and negotiated crosswinds flexibly to exploit both high altitude tailwinds and coastal soaring opportunities. We demonstrate that predicted airspeeds in heavy crosswinds depend strongly on orientation strategy and presumed preferred direction. Measured airspeeds increased with headwind and crosswind similarly to maximum‐range airspeeds based on full compensation for wind drift, yet remained ～ 30% lower than predicted by all strategies, resulting in slower and 30–35% costlier flight. Interestingly, more energy could be saved through adjustment of airspeed (median 40%) than via orientation strategy (median 4%). Therefore, despite remarkably flexible reaction to wind at sea, these gulls evidently minimized neither time nor energy expenditure. However, airspeeds were possibly over‐predicted by current aerodynamic models. This study emphasizes the importance of accounting for orientation strategy when assessing airspeed adjustments to wind and indicates that either the cost or adaptive ‘currency’ of extended flight among gulls may require revision.     

Dynamic body acceleration increases by 20% during flight ontogeny of greylag geese Anser anser

Despite our knowledge of the biophysical and behavioural changes during flight ontogeny in juvenile birds, little is known about the changes in the mechanical aspects of energy expenditure during early flight development, particularly in migratory species. Here, we investigate in a unique experimental setup how energy expended during flights changes over time beginning with early ontogeny. We calculate overall dynamic body acceleration (ODBA) as a proxy for energy expenditure in a group of hand raised greylag geese Anser anser trained to fly behind a microlight aircraft. We propose two potential hypotheses; energy expenditure either increases with increasing physiological suitability (the ‘physical development hypothesis’), or decreases as a result of behavioural improvements mitigating flight costs (the ‘behavioural development hypothesis’). There was a significant temporal increase of flight duration and ODBA over time, supporting the ‘physical development hypothesis’. This suggests that early on in flight ontogeny behavioural development leading to flight efficiency plays a weaker role in shaping ODBA changes than the increased physical ability to expend energy in flight. We discuss these findings and the implications of flight development on the life history of migratory species.     

How Cheap Is Soaring Flight in Raptors? A Preliminary Investigation in Freely-Flying Vultures

Measuring the costs of soaring, gliding and flapping flight in raptors is challenging, but essential for understanding their ecology. Among raptors, vultures are scavengers that have evolved highly efficient soaring-gliding flight techniques to minimize energy costs to find unpredictable food resources. Using electrocardiogram, GPS and accelerometer bio-loggers, we report the heart rate (HR) of captive griffon vultures (Gyps fulvus and G. himalayensis) trained for freely-flying. HR increased three-fold at take-off (characterized by prolonged flapping flight) and landing (>300 beats-per-minute, (bpm)) compared to baseline levels (80–100 bpm). However, within 10 minutes after the initial flapping phase, HR in soaring/gliding flight dropped to values similar to baseline levels, i.e. slightly lower than theoretically expected. However, the extremely rapid decrease in HR was unexpected, when compared with other marine gliders, such as albatrosses. Weather conditions influenced flight performance and HR was noticeably higher during cloudy compared to sunny conditions when prolonged soaring flight is made easier by thermal ascending air currents. Soaring as a cheap locomotory mode is a crucial adaptation for vultures who spend so long on the wing for wide-ranging movements to find food.     

To fly or not to fly: response of water striders to drying out of habitat

Abstract .1. Adults of Aquarius paludum inhabit both temporary and permanent water surfaces; Gerris latiabdominis lives only in temporary habitats. To clarify whether adults of both species stay in position or fly when habitats dry up, overwintered adults of A. paludum and G. latiabdominis collected in spring were reared under one of the following four conditions: (A) on water with sufficient food, (B) on water, starved, (C) on wet paper with sufficient food, (D) on wet paper, starved. All rearings were at LD 15.5:8.5 h, 20 ± 2 °C, resembling natural conditions for April to June.
2. Females of A. paludum in group C had lower fecundity than the control group A and some stopped laying eggs. When a water surface was restored, females that had entered reproductive diapause began to lay eggs again. In contrast, females of G. latiabdominis continued to lay eggs even when reared on damp paper.
3. Adults of A. paludum lived longer and adopted diapause posture with high frequency when starved and reared without a water surface. There were, however, no significant differences in the longevity or in the number of adults showing diapause posture between groups A and D of G. latiabdominis.
4. Females of A. paludum collected in the middle of May had more mature oocytes (mean: 20.8) than females of G. latiabdominis (mean: 8.0), and most had histolysed their indirect flight muscles; most females of G. latiabdominis had retained their flight muscles and flew readily.
5. When water surfaces dry, with food shortage, adults of A. paludum may survive in place for a relatively long time until the water surface returns. Adults of G. latiabdominis may fly to other water surfaces and reproduce without delay.     

Risk‐taking and the evolution of mechanisms for rapid escape from predators

Flight initiation distance (FID) is the distance at which an individual animal takes flight when approached by a human. This behavioural measure of risk‐taking reflects the risk of being captured by real predators, and it correlates with a range of life history traits, as expected if flight distance optimizes risk of predation. Given that FID provides information on risk of predation, we should expect that physiological and morphological mechanisms that facilitate flight and escape predict interspecific variation in flight distance. Haematocrit is a measure of packed red blood cell volume and as such indicates the oxygen transport ability and hence the flight muscle contracting reaction of an individual. Therefore, we predicted that species with short flight distances, that allow close proximity between a potential prey individual and a predator, would have high haematocrit. Furthermore, we predicted that species with large wing areas and hence relatively low costs of flight and species with large aspect ratios and hence high manoeuvrability would have evolved long flight speed. Consistent with these predictions, we found in a sample of 63 species of birds that species with long flight distances for their body size had low levels of haematocrit and large wing areas and aspect ratios. These findings provide evidence consistent with the evolution of risk‐taking behaviour being underpinned by physiological and morphological mechanisms that facilitate escape from predators and add to our understanding of predator–prey coevolution.     

The energy cost of flight: do small bats fly more cheaply than birds?

Flapping flight is one of the most expensive activities in terms of metabolic cost and this cost has previously been considered equal for the two extant vertebrate groups which evolved flapping flight. Owing to the difficulty of obtaining accurate measurements without disturbing flight performance, current estimates of flight cost within the group of small birds and bats differ by more than a factor of five for given body masses. To minimize the potential problem that flight behaviour may be affected by the measurements, we developed an indirect method of measuring flight energy expenditure based on time budget analysis in which small nectar-feeding bats (Glossophaginae) could continue their natural rhythm of flying and resting entirely undisturbed. Estimates of metabolic flight power based on 172 24-h time and energy budget measurements were obtained for nine individual bats from six species (mass 7–28 g). Metabolic flight power (P_F) of small bats was found to increase with body mass following the relation P_F = 50.2 M^0.771 (r² = 0.96, n = 13, P_F in W, M in kg). This is about 20–25% below the majority of current predictions of metabolic flight cost for small birds. Thus, either the flight cost of small birds is significantly lower than has previously been thought or, contrary to current opinion, small bats require less energy to fly than birds. Accepted: 29 September 1997     

Flight altitude of trans-Sahara migrants in autumn: a comparison of radar observations with predictions from meteorological conditions and water and energy balance models

Radar observations on the altitude of bird migration and altitudinal profiles of meteorological conditions over the Sahara desert are presented for the autumn migratory period. Migratory birds fly at an average altitude of 1016 m (a.s.l.) during the day and 571 m during the night. Weather data served to calculate flight range using two models: an energy model (EM) and an energy-and-water model (EWM). The EM assumes that fuel supply limits flight range whereas the EWM assumes that both fuel and water may limit flight range. Flight ranges estimated with the EM were generally longer than those with the EWM. This indicates that trans-Sahara migrants might have more problems balancing their water than their energy budget. However, if we assume fuel stores to consist of 70% instead of 100% fat (the remainder consisting of 9% protein and 21% water), predicted flight ranges of the EM and EWM largely overlap. Increased oxygen extraction, reduced flight costs, reduced exhaled air temperature, reduced cutaneous water loss and increased tolerance to water loss are potential physiological adaptations that would improve the water budget in migrants. Both the EM and EWM predict optimal flight altitudes in agreement with radar observations in autumn. Optimal flight altitudes are differently predicted by the EM and EWM for nocturnal spring migration. During spring, the EWM predicts moderately higher and the EM substantially higher flight altitudes than during autumn. EWM predictions are therefore in better agreement with radar observations on flight altitude of migrants over the Negev desert in spring than EM predictions.     

Perch-hunting in insectivorous Rhinolophus bats is related to the high energy costs of manoeuvring in flight

Foraging behaviour of bats is supposedly largely influenced by the high costs of flapping flight. Yet our understanding of flight energetics focuses mostly on continuous horizontal forward flight at intermediate speeds. Many bats, however, perform manoeuvring flights at suboptimal speeds when foraging. For example, members of the genus Rhinolophus hunt insects during short sallying flights from a perch. Such flights include many descents and ascents below minimum power speed and are therefore considered energetically more expensive than flying at intermediate speed. To test this idea, we quantified the energy costs of short manoeuvring flights (<2 min) using the Na-bicarbonate technique in two Rhinolophus species that differ in body mass but have similar wing shapes. First, we hypothesized that, similar to birds, energy costs of short flights should be higher than predicted by an equation derived for bats at intermediate speeds. Second, we predicted that R. mehelyi encounters higher flight costs than R. euryale, because of its higher wing loading. Although wing loading of R. mehelyi was only 20% larger than that of R. euryale, its flight costs (2.61 ± 0.75 W; mean ± 1 SD) exceeded that of R. euryale (1.71 ± 0.37 W) by 50%. Measured flight costs were higher than predicted for R. mehelyi, but not for R. euryale. We conclude that R. mehelyi face elevated energy costs during short manoeuvring flights due to high wing loading and thus may optimize foraging efficiency by energy-conserving perch-hunting.     

Why Do Kestrels Soar?

Individuals allocate considerable amounts of energy to movement, which ultimately affects their ability to survive and reproduce. Birds fly by flapping their wings, which is dependent on the chemical energy produced by muscle work, or use soaring-gliding flight, in which chemical energy is replaced with energy harvested from moving air masses, such as thermals. Flapping flight requires more energy than soaring-gliding flight, and this difference in the use of energy increases with body mass. However, soaring-gliding results in lower speeds than flapping, especially for small species. Birds therefore face a trade-off between energy and time costs when deciding which flight strategy to use. Raptors are a group of large birds that typically soar. As relatively light weight raptors, falcons can either soar on weak thermals or fly by flapping with low energy costs. In this paper, we study the flight behavior of the insectivorous lesser kestrel (Falco naumanni) during foraging trips and the influence of solar radiation, which we have adopted as a proxy for thermal formation, on kestrel flight variables. We tracked 35 individuals from two colonies using high frequency GPS-dataloggers over four consecutive breeding seasons. Contrary to expectations, kestrels relied heavily on thermal soaring when foraging, especially during periods of high solar radiation. This produced a circadian pattern in the kestrel flight strategy that led to a spatial segregation of foraging areas. Kestrels flapped towards foraging areas close to the colony when thermals were not available. However, as soon as thermals were formed, they soared on them towards foraging areas far from the colony, especially when they were surrounded by poor foraging habitats. This reduced the chick provisioning rate at the colony. Given that lesser kestrels have a preference for feeding on large insects, and considering the average distance they cover to capture them during foraging trips, to commute using flapping flight would result in a negative energy balance for the family group. Our results show that lesser kestrels prioritize saving energy when foraging, suggesting that kestrels are more energy than time-constrained during the breeding season.     

The physiology of life-history trade-offs: experimental analysis of a hormonally induced life-history trade-off in Gryllus assimilis

Abstract Adult Gryllus assimilis given an analog of juvenile hormone exhibited reduced flight muscles and enlarged ovaries similar to those found in naturally occurring flightless individuals of species that are polymorphic for dispersal capability. Control and hormone-treated (flightless) G. assimilis did not differ in the amount of food consumed or assimilated on any of three diets that differed in nutrient quantity. Thus, enhanced ovarian growth of flightless individuals resulted from increased allocation of internal nutrients to reproduction (i.e., a trade-off) rather than from increased acquisition of nutrients. Compared with flight-capable controls, flightless G. assimilis also had reduced whole-organism respiration, reduced respiration of flight muscles, and reduced lipid and triglyceride (flight fuel) reserves. These differences are remarkably similar to those between naturally occurring flightless and flight-capable morphs of other Gryllus species. Results collectively suggest that the increased allocation of nutrients to ovarian growth in flightless G. assimilis and other Gryllus species results from reduced energetic costs of flight muscle maintenance and/or the biosynthesis or acquisition of lipids. Reduction in these energetic costs appears to be an important driving force in the evolution of flightlessness in insects. Respiratory metabolism associated with flight capability utilizes an increasing proportion of the energy budget of crickets as the quantity of nutrients in the diet is decreased. This leads to a magnification of greater ovarian growth of flightless versus flight-capable individuals on nutrient-poor diets.     

Avoidance of headwinds or exploitation of ground effect—why do birds fly low?

ABSTRACT Birds often fly close to the ground or water. Wind shear theory predicts that wind speeds decline with proximity to the substratum, so birds might be expected to fly lower when flying upwind than when flying downwind. We tested this prediction and found that the wind shear equation is valid at heights below 4 m, with wind speed over a smooth surface ～40% lower at a height of 0.08 m than at 4 m. Birds that fly close enough to smooth substrata can also benefit energetically from ground effect, where vortices generated by their flight interact with the ground or water. This suggests that birds should use ground effect more when flying upwind than when flying downwind. We determined the percent time spent flying in ground effect by 21 species of passerine and non‐passerine birds flying in sheltered coastal aquatic and nearby terrestrial areas of County Cork, Ireland. We found that use of ground effect was uncommon for passerines, but common for a variety of non‐passerine waterbirds. However, phylogenetic analysis indicates no linkage between phylogeny and incidence of ground effect use and it is probable that incidence of use is determined by ecology rather than phylogeny. Great Cormorants (Phalacrocorax carbo) used ground effect most frequently over water (59.4% of time in flight). Over land, Barn Swallows (Hirundo rustica) used ground effect most often (19.8% of time). Phylogenetic contrasts regression analysis showed no significant relationship between use of ground effect and either wing aspect ratio or wing loading for 18 of our focal species, though simple linear regression analysis indicated that birds with greater wing loading used ground effect slightly (but significantly) more often. We found that 95% of Great Cormorants flying upwind used ground effect whereas only 35% did so when flying downwind. Few Black‐headed Gulls (Chroicocephalus ridibundus) used ground effect (probably because they fly high to locate prey), but still showed greater use when flying upwind (25%) than downwind (2.5%). When flying upwind in ground effect at the wind speeds encountered in our study, the velocity for minimum power (V_mp) for Great Cormorants was exceeded, suggesting theoretical benefits of about 14.3%. Our study indicates that several species exploit both wind shear and ground effect to minimize energy expenditure during commuting and foraging, but that others do not, because of either complexity of habitat morphology or the demands of their foraging ecology.     

Flying to feed or flying to mate: gender differences in the flight activity of tsetse (Glossina palpalis)

Abstract. To test the prediction that female and male tsetse should differ in their behaviour and the partitioning of their energy budgets to maximize their respective reproductive outputs, we investigated experimentally the relationship between blood intake, fat content and flight activity in virgin and mated female flies, following the same procedure as in a previous study on males.Those flies whose fat content was raised to higher levels by being fed more frequently performed more flight activity, but all females showed very little activity until day 4 after their last blood meal, thereby using only a small fraction of their fat reserves.This contrasted markedly with the large amount of flight performed by males on day 3, resulting in the depletion of their fat reserves.The difference is interpreted with respect to females flying only to find food approximately once every 3 days (and larviposition sites approximately once every 9 days), compared with males flying to find as many mates as possible during the earlier part of the feeding cycle when their energy reserves are high and feeding is a low priority.     

The energy costs of sexual dimorphism in mole-rats are morphological not behavioural

Different reproductive strategies of males and females may lead to the evolution of differences in their energetic costs of reproduction, overall energetic requirements and physiological performances. Sexual dimorphism is often associated with costly behaviours (e.g. large males might have a competitive advantage in fighting, which is energetically expensive). However, few studies of mammals have directly compared the energy costs of reproductive activities between sexes. We compared the daily energy expenditure (DEE) and resting metabolic rate (RMR) of males and females of two species of mole-rat, Bathyergus janetta and Georychus capensis (the former is sexually dimorphic in body size and the latter is not) during a period of intense digging when males seek females. We hypothesized that large body size might be indicative of greater digging or fighting capabilities, and hence greater mass-independent DEE values in males of the sexually dimorphic species. In contrast to this prediction, although absolute values of DEE were greater in B. janetta males, mass-independent values were not. No differences were apparent between sexes in G. capensis. By comparison, although RMR values were greater in B. janetta than G. capensis, no differences were apparent between the sexes for either species. The energy cost of dimorphism is most likely to be the cost of maintenance of a large body size, and not the cost of behaviours performed when an individual is large.     

Time‐restricted flight ability influences dispersal and colonization rates in a group of freshwater beetles

Variation in the ability to fly or not is a key mechanism for differences in local species occurrences. It is increasingly acknowledged that physiological or behavioral mechanisms rather than morphological differences may drive flight abilities. However, our knowledge on the seasonal variability and stressors creating nonmorphological differences in flight abilities and how it scales to local and regional occurrences is very limited particularly for small, short‐lived species such as insects. Here, we examine how flight ability might vary across seasons and between two closely related genera of freshwater beetles with similar geographical ranges, life histories, and dispersal‐related morphology. By combining flight experiments of >1,100 specimens with colonization rates in a metacommunity of 54 ponds in northern and eastern Europe, we have analyzed the relationship between flight ability and spatio‐environmental distribution of the study genera. We find profound differences in flight ability between the two study genera across seasons. High flight ability for Acilius (97% of the tested individuals flew during the experiments) and low for Graphoderus (14%) corresponded to the different colonization rates of newly created ponds. Within a 5‐year period, 81 and 31% of the study ponds were colonized by Acilius and Graphoderus, respectively. While Acilius dispersed throughout the season, flight activity in Graphoderus was restricted to stressed situations immediately after the emergence of adults. Regional colonization ability of Acilius was independent of spatial connectivity and mass effect from propagule sources. In contrast, Graphoderus species were closely related to high connectivity between ponds in the landscape. Our data suggest that different dispersal potential can account for different local occurrences of Acilius and Graphoderus. In general, our findings provide some of the first insights into the understanding of seasonal restrictions in flight patterns of aquatic beetles and their consequences for species distributions.     

Rain increases the energy cost of bat flight

Similar to insects, birds and pterosaurs, bats have evolved powered flight. But in contrast to other flying taxa, only bats are furry. Here, we asked whether flight is impaired when bat pelage and wing membranes get wet. We studied the metabolism of short flights in Carollia sowelli, a bat that is exposed to heavy and frequent rainfall in neotropical rainforests. We expected bats to encounter higher thermoregulatory costs, or to suffer from lowered aerodynamic properties when pelage and wing membranes catch moisture. Therefore, we predicted that wet bats face higher flight costs than dry ones. We quantified the flight metabolism in three treatments: dry bats, wet bats and no rain, wet bats and rain. Dry bats showed metabolic rates predicted by allometry. However, flight metabolism increased twofold when bats were wet, or when they were additionally exposed to rain. We conclude that bats may not avoid rain only because of sensory constraints imposed by raindrops on echolocation, but also because of energetic constraints.     

Fat is sexy for females but not males: the influence of body reserves on reproduction in snakes (Vipera aspis)

Reproduction is energetically expensive for both sexes, but the magnitude of expenditure and its relationship to reproductive success differ fundamentally between males and females. Males allocate relatively little to gamete production and, thus, can reproduce successfully with only minor energy investment. In contrast, females of many species experience high fecundity-independent costs of reproduction (such as migration to nesting sites), so they need to amass substantial energy reserves before initiating reproductive activity. Thus, we expect that the relationship between energy reserves and the intensity of reproductive behavior involves a threshold effect in females, but a gradual (or no) effect in males. We tested this prediction using captive vipers (Vipera aspis), dividing both males and females into groups of high versus low body condition. Snakes from each group were placed together and observed for reproductive behavior; sex-steroid levels were also measured. As predicted, females in below-average body condition had very low estradiol levels and did not show sexual receptivity, whereas males of all body condition indices had significant testosterone levels and displayed active courtship. Testosterone levels and courtship intensity increased gradually (i.e., no step function) with body condition in males, but high estradiol levels and sexual receptivity were seen only in females with body reserves above a critical threshold.     

Activity‐specific metabolic rates for diving,transiting, and resting at sea can be estimated from time–activity budgets in free‐ranging marine mammals

Time and energy are the two most important currencies in animal bioenergetics. How much time animals spend engaged in different activities with specific energetic costs ultimately defines their likelihood of surviving and successfully reproducing. However, it is extremely difficult to determine the energetic costs of independent activities for free‐ranging animals. In this study, we developed a new method to calculate activity‐specific metabolic rates, and applied it to female fur seals. We attached biologgers (that recorded GPS locations, depth profiles, and triaxial acceleration) to 12 northern (Callorhinus ursinus) and 13 Antarctic fur seals (Arctocephalus gazella), and used a hierarchical decision tree algorithm to determine time allocation between diving, transiting, resting, and performing slow movements at the surface (grooming, etc.). We concomitantly measured the total energy expenditure using the doubly‐labelled water method. We used a general least‐square model to establish the relationship between time–activity budgets and the total energy spent by each individual during their foraging trip to predict activity‐specific metabolic rates. Results show that both species allocated similar time to diving (~29%), transiting to and from their foraging grounds (~26–30%), and resting (~8–11%). However, Antarctic fur seals spent significantly more time grooming and moving slowly at the surface than northern fur seals (36% vs. 29%). Diving was the most expensive activity (~30 MJ/day if done non‐stop for 24 hr), followed by transiting at the surface (~21 MJ/day). Interestingly, metabolic rates were similar between species while on land or while slowly moving at the surface (~13 MJ/day). Overall, the average field metabolic rate was ~20 MJ/day (for all activities combined). The method we developed to calculate activity‐specific metabolic rates can be applied to terrestrial and marine species to determine the energetic costs of daily activities, as well as to predict the energetic consequences for animals forced to change their time allocations in response to environmental shifts.     

不同日龄和吊飞过程中桔小实蝇成虫飞行肌能量代谢相关酶活性的变化

【目的】明确桔小实蝇Bactrocera dorsalis(Hendel)飞行肌对能源物质的利用。【方法】通过生化方法测定了能源物质代谢相关5种酶[3-磷酸甘油醛脱氢酶(GAPDH)、3-磷酸甘油脱氢酶(GDH)、乳酸脱氢酶(LDH)、柠檬酸合酶(CS)和3-羟酰辅酶A脱氢酶(HOAD)]活性的变化。【结果】桔小实蝇成虫中所测的5种酶活性随日龄的变化而变化,4日龄GAPDH,GDH,LDH和CS活性最高,20日龄HOAD活性最高。吊飞过程中,GAPDH,GDH和CS的活性变化基本一致,随吊飞时间的延长活性逐渐升高;LDH和HOAD的活性变化雌、雄虫完全不同。雄虫LDH活性除吊飞2 h外其他时间均高于静息状态,雌虫则始终低于静息状态;雄虫HOAD活性只有吊飞24 h低于静息状态水平,而雌虫吊飞后HOAD活性一直在静息状态水平及以下波动。【结论】桔小实蝇飞行所利用的能源物质包括糖类和脂肪,以糖类能源为主。吊飞过程中,雄虫除可以进行高速有氧代谢以外,还具备一定的无氧代谢能力,而雌虫只进行有氧代谢;雄虫能利用脂肪供给能量,雌虫则几乎不动用脂肪。研究结果为进一步阐明桔小实蝇的迁飞行为机制提供了依据。