Size‐dependent insect flight energetics at different sugar supplies

Under most circumstances, large body size confers a higher fitness and is positively selected, whereas selection against large size is empirically poorly documented. Physiologically, according to the ¾ power law, larger animals have lower relative but higher absolute energy demands, such that large body size may become disadvantageous, particularly under fast locomotion in food‐limited environments. After a period of initial feeding on different sugar concentrations, we investigated size‐dependent energy content (reserves) at baseline and of females unflown (i.e. resting) or flown for 18 h in two (replicate) insect species: the yellow dung fly Scathophaga stercoraria and the yellow fever mosquito Aedes aegypti. Tethered adults of various sizes were tested in a flight mill. In both species, teneral glycogen, sugar, and lipid content increased with sugar availability, and isometrically or even hyper‐allometrically (slope > 1) with body size. Activity treatment also revealed the expected consumption effects. Both species increased their flight distance with sugar supply, although only larger mosquitoes flew longer. Crucially, larger females of both species disproportionately exhausted more glycogen and sugars (but not lipid) during flight. The mosquitoes appeared to adjust their flight more finely to their size‐dependent energy reserves at all sugar availabilities, whereas, in the dung flies, size‐dependent energy demands were detectable only with a low but not with an overly high sugar supply. Although we found a greater absolute and relative locomotory energy demand for the larger flies, which is in agreement with interspecific patterns in insects, this was (more than) compensated by their greater baseline energy reserves, resulting in the greater net flying capacity of larger individuals. Consequently, we found no evidence for energetic mechanisms limiting the performance of large flying insects under food limitation. The differences between the two species presumably relate to mosquitoes inherently being long distance flyers and dung flies being short distance flyers. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ●● , ●●–●●.     

Comparing the effects of five naturally occurring monosaccharide and oligosaccharide sugars on longevity and carbohydrate nutrient levels of a parasitic phorid fly, Pseudacteon tricuspis

Abstract.  The longevity and nutrient levels of Pseudacteon tricuspis provided with 1  m solutions of five naturally occurring sugars, fructose, glucose, sucrose, trehalose and melezitose, are compared. All but melezitose, result in significant increases in the longevity of P. tricuspis in comparison with sugar-starved flies (flies provided with water only). Sugar-starved female and male P. tricuspis have an average longevity of 3.3 and 4.1 days, respectively. Provision of free water in addition to sugar solution is necessary for optimum longevity by female and male flies. Longevity is increased by 2.4–2.6-fold by the two monosaccharides, fructose and glucose, and by 2.6–2.8-fold by the disaccharides, sucrose and trehalose. Phorid flies provided with the trisaccharide sugar, melezitose, had a marginal increase in lifespan (approximately 1 day), but this is not significantly different from the longevity of sugar-starved flies. Significantly greater levels of total sugars are detected in P. tricuspis fed the disaccharide sugars (sucrose, trehalose) or the monosaccharide sugars (fructose, glucose), compared with flies provided with melezitose (trisaccharide), or to sugar-starved flies. Fructose is not detected in sugar-starved flies, or in flies fed glucose or trehalose. However, high levels of fructose are detected in flies fed sucrose or fructose, whereas levels of fructose in melezitose-fed flies are intermediate. In general, significantly greater glycogen levels are detected in P. tricuspis fed sucrose, glucose, trehalose or fructose, compared with melezitose-fed or sugar-starved flies. Levels of total sugars and glycogen in sugar-fed flies are positively correlated with wing length, possibly indicating a higher accumulation of storage sugars by larger flies. These results are discussed in relation to the nutritional ecology of the phorid fly.     

Lifespan and patterns of accumulation and mobilization of nutrients in the sugar-fed phorid fly, Pseudacteon tricuspis

Abstract. The effect of sugar feeding on the survival of adult phorid fly Pseudacteon tricuspis is investigated. Flies fed 25% sucrose in aqueous solution continuously throughout their lifespan have greater longevity (mean ± SE longevity: female = 7.9 ± 0.8 days, male = 8.9 ± 0.9 days) than completely starved (provided no water and no sugar solution) flies, sugar-starved (provided water only) flies, or flies fed sugar solution only on their first day of adult life. Completely starved flies rarely lived beyond one day. Provision of water increases longevity by 2 days, and one full day of sugar feeding further increases longevity by an additional 1–2 days. Flies fed 50% sucrose have similar survivorship as those fed 25% sucrose. The temporal patterns of nutrient accumulation and utilization are also compared in P. tricuspis fed different diets: sugar-starved, sucrose-fed on the first day of adult life only, and sucrose-fed continuously. Adult P. tricuspis emerge with no gut sugars, and only minimal amounts of body sugars and glycogen. Although the levels of body sugars and glycogen decline gradually in sugar-starved flies, a single day of sugar feeding results in the accumulation of maximum amounts of gut sugars, body sugars and glycogen. High levels of these nutrients are maintained in female and male phorid flies fed on sucrose continuously over the observation period, whereas nutrient levels decline in flies fed only on the first day of life, beginning 1 day postfeeding. Female and male P. tricuspis emerge with an estimated 12.3 ± 2.3 and 7.2 ± 1 g of lipid reserves per fly, respectively. These teneral amounts represent the highest lipid levels detected in adult flies, irrespective of their diet, and are maintained over the life times of sucrose-fed female and male flies, but declined steadily in sugar-starved females. These data suggest that adult P. tricuspis are capable of converting dietary sucrose to body sugars and glycogen, but not lipids.     

Trade-off of energy metabolites as well as body color phenotypes for starvation and desiccation resistance in montane populations of Drosophila melanogaster

Storage of energy metabolites has been investigated in different sets of laboratory selected desiccation or starvation resistant lines but few studies have examined such changes in wild-caught populations of Drosophila melanogaster. In contrast to parallel selection of desiccation and starvation tolerance under laboratory selection experiments, opposite clines were observed in wild populations of D. melanogaster. If resistance to desiccation and starvation occurs in opposite directions under field conditions, we may expect a trade-off for energy metabolites but such correlated changes are largely unknown. We tested whether there is a trade-off for storage as well as actual utilization of carbohydrates (trehalose and glycogen), lipids and proteins in D. melanogaster populations collected from different altitudes (512-2500 m). For desiccation resistance, darker flies (> 50% body melanization) store more body water content and endure greater loss of water (higher dehydration tolerance) as compared to lighter flies (< 30% body melanization). Based on within population analysis, we found evidence for coadapted phenotypes i.e. darker flies store and actually utilize more carbohydrates to confer greater desiccation resistance. In contrast, higher starvation resistance in lighter flies is associated with storage and actual utilization of greater lipid amount. However, darker and lighter flies did not vary in the rate of utilization of carbohydrates under desiccation stress; and of lipids under starvation stress. Thus, we did not find support for the hypothesis that a lower rate of utilization of energy metabolites may contribute to greater stress resistance. Further, for increased desiccation resistance of darker flies, about two-third of total energy budget is provided by carbohydrates. By contrast, lighter flies derive about 66% of total energy content from lipids which sustain higher starvation tolerance. Our results support evolutionary trade-off for storage as well as utilization of energy metabolites for desiccation versus starvation resistance in D. melanogaster.     

Life history evolution and cellular mechanisms associated with increased size in high‐altitude Drosophila

Understanding the physiological and genetic basis of growth and body size variation has wide‐ranging implications, from cancer and metabolic disease to the genetics of complex traits. We examined the evolution of body and wing size in high‐altitude Drosophila melanogaster from Ethiopia, flies with larger size than any previously known population. Specifically, we sought to identify life history characteristics and cellular mechanisms that may have facilitated size evolution. We found that the large‐bodied Ethiopian flies laid significantly fewer but larger eggs relative to lowland, smaller‐bodied Zambian flies. The highland flies were found to achieve larger size in a similar developmental period, potentially aided by a reproductive strategy favoring greater provisioning of fewer offspring. At the cellular level, cell proliferation was a strong contributor to wing size evolution, but both thorax and wing size increases involved important changes in cell size. Nuclear size measurements were consistent with elevated somatic ploidy as an important mechanism of body size evolution. We discuss the significance of these results for the genetic basis of evolutionary changes in body and wing size in Ethiopian D. melanogaster.     

Does gliding when pregnant select for larger females?

For terrestrial vertebrates, gliding imposes unique constraints on the interaction of body mass and structural size, particularly with reference to minimizing wing loading. Females of gliding animals experience increases in wing loading during pregnancy or gravidity, and selection may favour increased structural size to compensate for the added mass. We tested whether pregnant southern flying squirrels Glaucomys volans had similar wing loading as males, and whether females with lower wing loading bore heavier litters, than those with greater wing loading. Males had greater wing loading than females, regardless of the latter's reproductive state (males: 38.4±3.62 N m⁻², pregnant females: 30.7±4.21 N m⁻² and non-pregnant females: 26.8±5.13 N m⁻²). The slope of the linear relationship between planar surface area and body mass was similar between pregnant females and males, however ( F =0.383, P =0.322). Thus female flying squirrels may optimize their litter mass to minimize wing loading during pregnancy. Contrary to our prediction, females with greater wing loading had heavier litters than those with lower wing loading, which suggests reproductive output may be influenced by other ecological factors.     

Effect of body size on energy reserves in Culex pipiens pallens females (Diptera: Culicidae)

We examined the relationship between body size and energy reserves of female Culex pipiens pallens under laboratory conditions. Small and large‐sized adult mosquitoes were obtained from larvae reared at different densities. There was a significantly positive relationship between pupal body weight and adult wing length depending on body sizes. Studies showed large females contained significantly higher contents of glycogen and lipid but lower protein stores in comparison to small females. Large sized mosquitoes accumulated higher percentages of glycogen (21.2% vs 15.9%) and lipid (45.8% vs 30.7%) but lower levels of protein (33.0% vs 53.4%) than small females. This result suggests that larval‐derived glycogen and lipid stores may be important in determining the body size of female Cx. pipiens pallens.     

Sexual size dimorphism and sex ratios in dragonflies (Odonata)

Sexual size dimorphism and biased sex ratios are common in animals. Rensch's rule states that sexual size dimorphism (SSD) would increase with body size in taxa where males are larger than females and decrease with body size in taxa where females are larger. We tested this trend in dragonflies (Odonata) by analysing body size of 21 species and found support for Rensch's rule. The increase in SSD with increasing size among species can be explained by sexual selection favouring large males. We also estimated the slope of the relationship between sex ratio and size ratio in populations of the 21 species. A negative slope would suggest that the larger sex suffers from high mortality in the larval stage, consistent with riskier foraging. The slope of this relationship was negative, but after correcting for phylogentic non-independence with independent contrasts the relationship was no longer statistically significant, perhaps because of phylogenic inertia or low sample size.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 507–513.     

Recent increase in body size of the American marten Martes americana in Alaska

Using museum specimens, we studied recent changes in skull size of the American marten Martes americana , in continental Alaska. In Alaska, global warming has resulted in milder winters that may contribute to an improved food supply in the wild. In the present study, we tested the hypothesis that body size of the marten had increased during the second half of the 20th century, in response to global warming. We found that skull size, and by implication body size, increased significantly during the second half of the 20th century, possibly due to an improved food supply and/or lower metabolic demands in winter. Improved food availability in winter may result from the improved nutritional conditions for prey, and/or from increased access to prey resulting from a longer snow-free season. Longitude had a significant positive effect on skull size and a significant negative effect on teeth size. In Alaska, the climate is milder along the western coast and becomes harsher inland. Hence, the milder climate was associated with larger body size providing further support for our prediction that body size of the American marten was influenced by food availability and reduced energy expenditure. The negative relationship between longitude and teeth size may indicate a trend towards a larger prey in inland marten populations, but we have no data to support or refute this hypothesis.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 701–707.     

Biochemical content, energy composition and reproductive effort in the broadcasting sea star Asterias vulgaris over the spawning period

To assess sex differences in reproductive effort, we examined the biochemical composition and energetic content of the principal body components of the broadcast spawning sea star Asterias vulgaris in the Mingan Islands in the northern Gulf of St. Lawrence, eastern Canada. The body wall was the most stable body component, showing no variations in mass or in lipid and protein content (and total energetic content) between sexes or during spawning. Patterns in the gonads differed between sexes and with spawning. The lipid, protein and carbohydrate content of the ovary dropped during spawning, while only the protein content of the testis decreased significantly. Reproductive effort, expressed as loss of energy in the gonads during spawning for an individual weighing 10 g in underwater mass (8.2 cm in radius), was six times greater in females (49.5 kJ) than males (7.9 kJ). The energetic content of the pyloric caeca also decreased during spawning, by 17.7 kJ in females and 21.5 kJ in males, mainly due to a decrease in lipids. If this decrease is included as reproductive effort, it lessens the gender difference. The caecum decrease possibly represented expenditures due to formation of aggregations or the expulsion of gametes during spawning. Effectively, we observed aggregations during a massive spawning in this population. The sex ratio did not differ from 1:1 in all size classes sampled. This suggests that, unless males suffer higher mortality, females manage to allocate as much energy to somatic growth as males, possibly by feeding at higher rates to compensate for their higher reproductive effort. Stomach protein content tended to be higher in females than males and may indicate greater muscular development to facilitate digestion.     

Annual variation in energy reserves in male three-spined stickleback, Gasterosteus aculeatm L. (Pisces, Gasterosteidae)

Wet and dry weights of tissue were measured and concentrations of glycogen, lipid and protein were estimated for the liver, gonad and carcass of male sticklebacks from an annual population collected each month over one complete year. In young-of-the-year there is one period of rapid weight gain, in all three body regions (liver, carcass and gonads) but particularly of the carcass, in the autumn and a second in spring and early summer. This is accompanied by an increase in the water content of all three body regions. The gonadosomatic index also increases sharply in spring and early summer. Young sticklebacks accumulate lipid and glycogen slowly during the autumn and winter of their first year of life and more rapidly from late winter to early summer. Thus, the period of most rapid accumulation of these reserves coincides with the time when body weight and gonad maturation are also increasing sharply. Lipid and glycogen levels fall during the reproductive season in those males that breed, so that by July they are reduced to 43% and 37% (respectively) of their peak values in May. Levels of protein increase throughout the year as the fish grow, but in breeding males by July the concentration of protein in the carcass falls to 70% of pre-breeding levels. Breeding males therefore reach the end of the reproductive season with their total energy reserves severely reduced, and consequently they suffer a very high mortality. In contrast, adult males that fail to reproduce survive beyond the breeding season. They continue to gain weight and to accumulate lipid and glycogen from August to September, but these energy reserves fall (to levels comparable to those of post-breeding fish) in December, when these fish probably die. These results demonstrate that in male sticklebacks, growth and gonad maturation can be sustained in parallel with the accumulation of energy reserves, which are subsequently extensively depleted as a result of reproductive activities.     

Selection for increased maternal body volume does not differ between two Scincella lizards with different reproductive modes

Body size and shape are among the most important determinants of reproductive output in diverse animal taxa. We compared morphology and reproductive output between two Scincella lizards (Scincidae), Scincella modesta (oviparous) and Scincella reevesii (viviparous), to examine whether viviparous females compensate for their lower reproductive output by modifying maternal body size and/or shape. As predicted, reproductive output was lower in S. reevesii than in S. modesta when corrected for body size. The two lizards differed morphologically, but were similar in three aspects: females were the larger sex, the relative head size was greater in adult males, and the relative abdomen size was greater in adult females. Sexual dimorphism in abdomen length (AL) was more evident in S. reevesii than in S. modesta, but this difference was attributable to a smaller sexual difference in AL in S. modesta, rather than to the greater relative maternal AL in S. reevesii. Female S. reevesii on average were larger than female S. modesta in snout-vent length (SVL), but this increase in overall body size cannot be viewed as a way of allowing female S. reevesii to compensate for lower reproductive output, as the linear slope of reproductive output against maternal SVL did not differ between the two species. Our data show that selection for increased maternal body volume does not differ between the two Scincella lizards with different reproductive modes.     

Sublethal effect of brood parasitism on the grass-carrying wasp Isodontia mexicana

Abstract.  1. The nests of solitary, nest-provisioning wasps (Sphecidae and Crabronidae) are commonly attacked by brood parasites, including flies of the families Phoridae and Sarcophagidae. Larvae of the flies commonly kill the wasp offspring directly or starve it by consuming prey provided by the adult female wasps.
2. To determine whether brood parasitic flies can have a sublethal effect (i.e. reduced body size) on wasp offspring, nests of the wasp Isodontia mexicana were collected at two field sites in upstate New York, U.S.A. Nest diameter had no effect on the probability that a wasp cell would be inhabited by a brood parasite. Most offspring that developed in cells also containing phorids or sarcophagids managed to complete development and emerge as adults. Nevertheless, they had significantly smaller body size than conspecifics emerging from unparasitised cells in which the developing wasp did not have to compete for food (which in this species consists of tree crickets and katydids). Apparently, this is the first time that a sublethal effect of brood parasitism on offspring body size has been quantified for a solitary wasp species. Known effects of body size on the reproductive success of adult wasps suggest that sublethal consequences of the presence of brood parasites may have a significant effect on the fitness of adult I. mexicana .     

Measuring temporal variation in reproductive output reveals optimal resource allocation to reproduction in the northern grass lizard, Takydromus septentrionalis

We measured the reproductive output of Takydromus septentrionalis collected over 5 years between 1997 and 2005 to test the hypothesis that reproductive females should allocate an optimal fraction of accessible resources in a particular clutch and to individual eggs. Females laid 1–7 clutches per breeding season, with large females producing more, as well as larger clutches, than did small females. Clutch size, clutch mass, annual fecundity, and annual reproductive output were all positively related to female size (snout–vent length). Females switched from producing more, but smaller eggs in the first clutch to fewer, but larger eggs in the subsequent clutches. The mass-specific clutch mass was greater in the first clutch than in the subsequent clutches, but it did not differ among the subsequent clutches. Post-oviposition body mass, clutch size, and egg size showed differing degrees of annual variation, but clutch mass of either the first or the second clutch remained unchanged across the sampling years. The regression line describing the size–number trade-off was higher in the subsequent clutch than in the first clutch, but neither the line for first clutch, nor the line for the second clutch varied among years. Reproduction retarded growth more markedly in small females than in large ones. Our data show that: (1) trade-offs between size and number of eggs and between reproduction and growth (and thus, future reproduction) are evident in T. septentrionalis ; (2) females allocate an optimal fraction of accessible resources in current reproduction and to individual eggs; and (3) seasonal shifts in reproductive output and egg size are determined ultimately by natural selection.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 315–324.     

Resource availability modulates the effect of body size on reproductive development

Within-species variation in animal body size predicts major differences in life history, for example, in reproductive development, fecundity, and even longevity. Purely from an energetic perspective, large size could entail larger energy reserves, fuelling different life functions, such as reproduction and survival (the “energy reserve” hypothesis). Conversely, larger body size could demand more energy for maintenance, and larger individuals might do worse in reproduction and survival under resource shortage (the “energy demand” hypothesis). Disentangling these alternative hypotheses is difficult because large size often correlates with better resource availability during growth, which could mask direct effects of body size on fitness traits. Here, we used experimental body size manipulation in the freshwater cnidarian Hydra oligactis, coupled with manipulation of resource (food) availability to separate direct effects of body size from resource availability on fitness traits (sexual development time, fecundity, and survival). We found significant interaction between body size and food availability in sexual development time in both males and females, such that large individuals responded less strongly to variation in resource availability. These results are consistent with an energy reserve effect of large size in Hydra. Surprisingly, the response was different in males and females: small and starved females delayed their reproduction, while small and starved males developed reproductive organs faster. In case of fecundity and survival, both size and food availability had significant effects, but we detected no interaction between them. Our observations suggest that in Hydra, small individuals are sensitive to fluctuations in resource availability, but these small individuals are able to adjust their reproductive development to maintain fitness.     

An optimum body size for mammals? Comparative evidence from bats

1. The distribution of body sizes among mammalian species has been modelled by Brown, Marquet & Taper (1993), who suggest that reproductive power (the rate at which energy from the environment is channelled into offspring production) is maximized at a size of 100 g, and the observed size distribution among species reflects the way reproductive power depends on size. The model makes a testable prediction about life-history allometries: namely, that components of reproductive power should not scale linearly with body size but should change sign at the optimum size.
2. A large set of life-history data from a single clade of small mammals, the bats (Order: Chiroptera), was analysed to test this key prediction. The analyses in this study offer no support for the idea that allometries of reproductive power change sign in bats, either at 100 g or at any other size. Furthermore the life-history allometries of bats, which are mostly below the 100 g optimum, were broadly the same as in mammalian taxa larger than the optimum size.
3. These findings together contradict a key prediction of Brown et al. 's (1993) model to explain the skewed body size distribution across mammalian species.     

Sex-specific differences in desiccation resistance and the use of energy metabolites as osmolytes in Drosophila melanogaster flies acclimated to dehydration stress

In the Indian subcontinent, there are significant between-population variations in desiccation resistance in Drosophila melanogaster, but the physiological basis of adult acclimation responses to ecologically relevant humidity conditions is largely unknown. We tested the hypothesis that increased desiccation resistance in acclimated flies is associated with changes in cuticular permeability and/or content of energy metabolites that act as osmolytes. Under an ecologically relevant humidity regime (~50 % relative humidity), both sexes showed desiccation acclimation which persisted for 2–3 days. However, only females responded to acclimation at ~5 % relative humidity (RH). Acclimated flies exhibited no changes in the rate of water loss, which is consistent with a lack of plastic changes in cuticular traits (body melanization, epicuticular lipid). Therefore, changes in cuticular permeability are unlikely in drought-acclimated adult flies of D. melanogaster. In acclimated flies, we found sex differences in changes in the content of osmolytes (trehalose in females versus glycogen in males). These sex-specific changes in osmolytes are rapid and reversible and match to corresponding changes in the increased desiccation resistance levels of acclimated flies. Further, the increased content of trehalose in females and glycogen in males support the bound-water hypothesis for water retention in acclimated flies. Thus, drought acclimation in adult flies of D. melanogaster involves inducible changes in osmolytes (trehalose and glycogen), while there is little support for changes in cuticular permeability.     

Effects of energetic reserves on behavioral patterns of Mediterranean fruit flies (Diptera: Tephritidae)

The objective of this study was to associate levels of nutritional reserves (specifically lipids, sugars, and glycogen) in individual Mediterranean fruit flies, Ceratitis capitata (Diptera: Tephritidae), with observed patterns of behavior in the field. We collected females (n=255) and males (n=181) from the field, recording the time of collection and the activity they were engaged in when collected. Subsequently, we employed colorimetric biochemical techniques to determine the precise amounts of lipids, sugars, and glycogen in each individual. Lipid and sugar levels in males varied significantly according to the time of collection and the type of activity. Lipid and sugar levels in females did not vary in this manner. Sugar levels in both males and females were highest during the evening, when most feeding occurs. Males that engaged in sexual signaling in leks during the mid-afternoon had relatively low sugar and high lipid levels. Males engaged in the alternative mating tactic of fruit guarding had relatively high sugar and low lipid contents. Glycogen levels in males were high in the mornings, and a decline in glycogen content was associated with participation in leks; however, female glycogen levels did not vary significantly with time of day or activity. Our results provide quantitative evidence for the role nutrient reserves play in driving patterns of male reproductive behavior, yet suggest that factors other than sugar and lipid reserves constrain female behavior. Received: 8 April 1997 / Accepted: 23 June 1997     

The effect of exercise and captivity on energy partitioning, reproductive maturation and fertilization success in adult sockeye salmon

Releasing a population of adult Fraser River sockeye salmon Oncorhynchus nerka from an energetically more expensive non‐feeding natural migration (1089 km, natal groups) and allowing them to sexually mature in either a moderate flow (captive exercise group) or a no flow (captive non‐exercise group) environment resulted in significant differences in body energy densities (MJ kg⁻¹) among groups (ranked: initial natural migrator > non‐exercise > exercise > natal arrivals). Similarly, per cent body lipids were significantly lower in exercise females compared with non‐exercise females, experimentally confirming the central role lipid catabolism has in powering swimming. Although restricting exercise did result in greater body energy reserves at spawning, this did not result in a reallocation of energy to reproduction, as fecundity, egg size and gonad composition were unchanged among the groups. Furthermore, non‐exercise females had delayed maturity, lower egg deposition rates, and were more likely to die prior to egg ovulation than exercise females and natal spawners. Eggs from captive exercise adult females were more likely to survive to the eyed stage than eggs from captive non‐exercise females, but both captive groups had significantly lower egg fertilization success than natal spawners suggesting that confinement stress played a role in some of results presented. There were no significant differences in plasma levels of glucose, lactate, cortisol and reproductive hormones among the groups, but correlations among acute and chronic indicators of stress were significant among individuals. These results indicate that exercise during the late stages of maturation, as might occur normally, may be required for optimizing reproductive maturation and maximizing reproductive success.