Energetic costs, underlying resource allocation patterns, and adaptive value of predator-induced life-history shifts

We studied costs and benefits of life history shifts of water fleas (genus Daphnia ) in response to infochemicals from planktivorous fish. We applied a dynamic energy budget model to investigate the resource allocation patterns underlying the observed life history shifts and their adaptive value under size selective predation in one coherent analysis. Using a published data set of life history shifts in response to fish infochemicals we show that Daphnia invests less energy in somatic growth in the fish treatment. This observation complies with theoretical predictions on optimal resource allocation. However, the observed patterns of phenotypic plasticity cannot be explained by changes in resource allocation patterns alone because our model-based analysis of the empirical data clearly identified additional bioenergetic costs in the fish treatments. Consequently, the response to fish kairomone only becomes adaptive if the intensity of size selective predation surpasses a certain critical level. We believe that this is the first study that puts resource allocation, energetic costs, and adaptive value of predator induced life-history shifts – using empirical data – into one theoretical framework.     

Experimental testing of dynamic energy budget models

1. Dynamic energy budget (DEB) models describing the allocation of assimilate to the competing processes of growth, reproduction and maintenance in individual organisms have been applied to a variety of species with some success. There are two contrasting model formulations based on dynamic allocation rules that have been widely used (net production and net assimilation formulations). However, the predictions of these two classes of DEB models are not easily distinguished on the basis of simple growth and fecundity data.
2. It is shown that different assumptions incorporated in the rules determining allocation to growth and reproduction in two classes of commonly applied DEB models predict qualitatively distinct patterns for an easily measured variable, cumulative reproduction by the time an individual reaches an arbitrary size.
3. A comparison with experimental data from Daphnia pulex reveals that, in their simplest form, neither model predicts the observed qualitative pattern of reproduction, despite the fact that both formulations capture basic growth features.
4. An examination of more elaborate versions of the two models, in which the allocation rules are modified to account for brief periods of starvation experienced in the laboratory cultures, reveals that a version of the net production model can predict the qualitative pattern seen for cumulative eggs as a function of mass in D. pulex . The analysis leads to new predictions which can be easily tested with further laboratory experiments.     

Egg size evolution and energetic constraints on population dynamics.

We use population models that are based on dynamic energy budget models for individuals in order to study the evolution of offspring size and its relationship to the evolution of population dynamics. We show the existence of alternative evolutionarily stable strategies for offspring investment strategy resulting from a trade off between offspring number and time-to-maturity. The model predicts egg energy in Daphnia magna well, and suggests that the observed egg energy in D. magna is the result of selection for minimal egg investment constrained by minimum viable egg energy, combined with selection for a juvenile energy reserve. The selection for minimal egg size pushes populations toward chaotic dynamics. However, the minimum viable egg size combined with low efficiency of conversion of energy to new biomass is sufficient to keep population dynamics out of chaos.     

Does body storage act as a food-availability cue for adaptive adjustment of egg size and number in Daphnia magna?

1. The hypothesis that body storage is used by daphniids as a physiological cue for adaptively adjusting egg size and number to food availability was tested.
2. Egg size and number were examined to see whether they are related to individual variation in body storage independent of maternal size, genotype and food level. Egg mass and brood size (number of eggs in a brood) were compared to somatic mass, all adjusted to maternal body length, at two food levels in two parthenogenetic clones of Daphnia magna .
3. The prediction that adjusted brood size should increase with body storage, whereas adjusted egg mass should decrease with increasing body storage, was not fulfilled as seven of eight comparisons failed to fit the expectation.
4. It is concluded that body storage is probably not the food‐availability cue used by daphniids to control intrabrood resource allocation. Other possibilities, such as chemical cues emitted by food organisms and by coexisting cladocerans, are briefly discussed.     

Energy allocation in the cladoceran Daphnia magna Straus,under starvation and refeeding

Summary Models incorporating the energetics of individual daphnids (Cladocera) have been developed to predict the effect of environmental variables, particularly food availability, on population dynamics. One of them, that of Kooijman (1986), assumes that all assimilated energy enters a storage compartment prior to use in production and metabolism, and that under starvation the stores are used to support maintenance, reproduction and somatic growth, in that order of priority. This predicts that, under starvation, reproduction and growth will continue for a time, and that after they cease death will be immediate. Another model, that of McCauley et al. (1990), assumes that assimilated energy is used directly for maintenance and production, and that stores are accumulated to support maintenance metabolism under starvation. This predicts that growth and reproduction should cease immediately upon starvation and that death will not be immediate. We have carried out laboratory experiments, manipulating starvation time, on Daphnia magna to distinguish between these two models. The results support features of both models in that reproduction, but not growth, ceases upon starvation. We therefore developed a third model in which both maintenance and growth are supported from stores under starvation, with maintenance taking priority over growth under these conditions.     

The causes and consequences of variation in offspring size: a case study using Daphnia

Offspring size can have large and direct fitness implications, but we still do not have a complete understanding of what causes offspring size to vary. Daphnia (water fleas) generally produce fewer and larger offspring when food is limited. Here, we use a mathematical model to show that this could be explained by either: (1) an advantage of producing larger eggs when food is limited; or (2) a lower boundary on egg volume (below which eggs do not have sufficient resources to be viable), that is similar in volume to the evolutionarily stable egg volume predicted by standard clutch size models. We tested the first possibilities experimentally by placing offspring from mothers kept at two food treatments (high and low - leading to relatively small and large eggs respectively) into two food treatments (same as maternal treatments, in a fully factorial design) and measuring their fitness (reproduction, age at maturity, and size at maturity). We also tested survival under starvation conditions of offspring produced from mothers at low and high food treatments. We found that (larger) offspring produced by low-food mothers actually had lower fitness as they took longer to reproduce, regardless of their current food treatment. Additionally, we found no survival advantage to being born of a food-stressed mother. Consequently, our results do not support the hypothesis that there is an advantage to producing larger eggs when food is limited. In contrast, data from the literature support the importance of a lower boundary on egg size.     

Environmental stress and reproduction in Drosophila melanogaster : starvation resistance, ovariole numbers and early age egg production

Background  

The Y model of resource allocation predicts a tradeoff between reproduction and survival. Environmental stress could affect a tradeoff between reproduction and survival, but the physiological mechanisms underlying environmental mediation of the tradeoff are largely unknown. One example is the tradeoff between starvation resistance and early fecundity. One goal of the present study was to determine if reduced early age fecundity was indeed a robust indirect response to selection for starvation resistance, by investigation of a set of D. melanogaster starvation selected lines which had not previously been characterized for age specific egg production. Another goal of the present study was to investigate a possible relationship between ovariole number and starvation resistance. Ovariole number is correlated with maximum daily fecundity in outbred D. melanogaster. Thus, one might expect that a negative genetic correlation between starvation resistance and early fecundity would be accompanied by a decrease in ovariole number.     

Parental care trade‐offs in the inter‐brood phase in Barn Swallows Hirundo rustica

In seasonal environments with limited time and energy resources, double‐brooded birds face trade‐offs in the timing of their two reproductive attempts and in the effort allocated to the first and the second broods. In the Barn Swallow Hirundo rustica a long care period for the first brood enhances the survival of first‐brood chicks, but also delays the start of the second brood, which in turn reduces the survival prospects of second‐brood chicks. Probably as a response to this trade‐off, double‐brooded Barn Swallows reduce the period of post‐fledging care for first‐brood fledglings. By radiotracking whole families, we investigated the determinants of this behaviour and its consequences for the survival of the first‐brood fledglings. The end of the females’ investment in post‐fledging care of the first brood was related to the beginning of egg synthesis for the second clutch. With the start of egg synthesis, females significantly reduced provisioning rates to the first‐brood fledglings to less than one‐fifth of the previous rates, while the proportion of time they spent foraging remained high. Assuming that the females’ foraging success was constant, we conclude that their energy income was allocated to egg production rather than fledgling provision. Males did not compensate for the females’ reduced feeding rates. Thus the start of egg production for the second clutch had a marked effect on the quantity of food received by first‐brood fledglings. In parallel with the changes in parental behaviour and provisioning rates, we observed a marked drop in the daily survival rate of first‐brood chicks. These results support the hypothesis that females face a strong trade‐off in the allocation of energy to subsequent broods. Energy allocation to a second clutch involves a cost in terms of reduced provisioning, and as a result the survival of first‐brood chicks is compromised. This is probably outweighed by the improved success of an early second brood.     

Phenotypic plasticity and constancy of life-history traits in laboratory clones of Daphnia magna Straus: effects of neonatal length

1. The phenotypic constancy of four laboratory Daphnia magna clones in fitness-related life-history traits, such as age and clutch size at maturity, was studied among consecutive experimental runs in differing food environments.
2. A significant part of the observed clonal and genetic-by-environmental variation in age and clutch size at maturity was explained by experimentally uncontrollable variations in neonatal body length.
3. Despite food availability, neonatal length determined the number of instars invested to maturity and thus maturation age. Clonal differences in neonatal length and thus in maturation instar occurrence across environments explained most of the clonal variability observed in maturation age.
4. Although interclonal differences in clutch size existed, most of the phenotypic plasticity observed for clutch size was mediated by clonal differences in neonatal length.
5. Clonal differences in neonatal length and in the occurrence of maturation instars across environments dramatically affected the body length of instar IM-2 where provisioning of eggs take place. Since clutch size is determined from clutch mass and clutch mass was strongly related to the body length of instar IM-2, clonal differences across environments in body length of instar IM-2 mirrored clonal differences across environments in clutch size.
6. The results reported in the present study show that maternally mediated traits such as neonatal length affect how genotypes respond to different environmental selection regimes (genetic-by-environmental interaction). Future research needs to focus on the effects of neonatal length on the heritability or genetic variation of the reaction norms, since prediction of the response to selection is a key research objective in quantitative genetic studies.     

The trade‐off between clutch size and egg mass in tree swallows Tachycineta bicolor is modulated by female body mass

Egg production is a costly component of reproduction for female birds in terms of energy expenditure and maternal investment. Because resources are typically limited, clutch size and egg mass are expected to be constrained, and this putative trade‐off between offspring number and size is at the core of life history theory. Nevertheless, empirical evidence for this trade‐off is equivocal at best, as individual heterogeneity in resource acquisition and allocation may hamper the detection of the negative correlation between egg number and mass within populations. Here, we investigated how female body mass and landscape composition influences clutch size, egg mass, and the relationship between these two traits. To do so, we fitted linear mixed models using data from tree swallows Tachycineta bicolor breeding in a network of 400 nestboxes located along a gradient of agricultural intensity between 2004 and 2011. Our dataset comprised 1463 broods for clutch size analyses and 4371 eggs (from 847 broods laid between 2005–2008) for egg mass analyses. Our results showed that agricultural intensity negatively impacted clutch size, but not egg mass nor the relationship between these two traits. Female mass, on the other hand, modulated the trade‐off between clutch size and egg mass. For heavier females, both traits increased jointly, without evidence of a trade‐off. However, for lighter females, there was a clear negative relationship between clutch size and egg mass. This work shows that accounting for individual heterogeneity in body mass allows the detection of a clutch size/egg mass trade‐off that would have remained undetected otherwise. Identifying habitat and individual effects on resource allocation towards reproductive traits may help bridging the gap between predictions from theory and empirical evidence on life history trade‐offs.     

Maternal effects in Daphnia: what mothers are telling their offspring and do they listen?

Maternal effects can significantly impact offspring performance. Provisioning of offspring with energy stores can quantitatively alter their growth rates, survivorship, and future fecundity, and influence population regulatory mechanisms. In this paper, we show that maternal effects can also qualitatively affect offspring reproduction (i.e. their mode of reproduction). The freshwater herbivore Daphnia pulex can change the amount of energy allocated between asexual and ephippial egg production. Our experiments on individuals, experiencing “step‐up” or “step‐down” food manipulations, reveal that offspring qualitatively shift their energy allocation away from asexual reproduction to ephippial egg production when there is a simple mismatch between maternal and offspring food environments. We show that the response is asymmetric with respect to changes in food level, ephippial egg production is higher with a greater mismatch between environments, and that the effect can be observed in dynamic experimental populations. These results point to a “generational memory” that could challenge our interpretation of field patterns and mechanisms influencing population dynamics in Daphnia–algal systems.     

Juvenoid hormone methyl farnesoate is a sex determinant in the crustacean Daphnia magna

Daphnids (Daphnia magna) utilize cyclic parthenogenesis as a reproductive strategy. During periods of abundant resources, these organisms reproduce asexually. In response to environmental cues that signal the onset of environmental adversity, daphnids produce males and reproduce sexually. The environmental cues that stimulate the sexual reproductive phase are well known; however, the endocrine signals that transduce these environmental cues remain unknown. The present study was undertaken to test the hypothesis that the crustacean juvenoid hormone, methyl farnesoate, is a male sex determinant in this species. Continuous exposure to aqueous concentrations of methyl farnesoate greater than approximately 30 nM stimulated a concentration-dependent production of male-containing broods of organisms. Short-term exposures to methyl farnesoate during periods of egg and embryo maturation revealed that male sex determination occurred during a specific 12-hour period of ovarian egg development. Exposure of eggs to 400 nM methyl farnesoate during this sensitive developmental period resulted in the production of all-male broods of offspring, while exposure to concentrations as low as 52 nM produced mixed broods of males and females. This active concentration range of methyl farnesoate is consistent with levels measured in the hemolymph of some decapod crustaceans. These results demonstrate that methyl farnesoate is capable of programming daphnid embryos to develop into males and is likely the endocrine factor responsible for initiating the sexual reproductive phase in these organisms.     

Butterfly flight activity affects reproductive performance and longevity relative to landscape structure

Due to an overlap in the resources used by the flight muscles with the resources used during egg production, it has been hypothesised that an increased dispersal in fragmented landscapes may result in a physiological trade-off between flight and reproduction. In a common garden experiment, we investigated the effects of increased flight on the reproductive output of female speckled wood butterflies (Pararge aegeria, L.) from closed continuous woodland populations versus open highly fragmented agricultural landscapes in central France. Our flight treatment significantly affected resource allocation to egg size, but had no effect on mean daily fecundity. This treatment effect was similar for females from the two landscapes of origin, and suggests that energetic costs associated with increased flight result in a decrease in resource allocation to egg provisioning. There was a landscape-specific effect of flight on longevity: flight reduced longevity by 21% in woodland females, but had no affect on agricultural females. This result suggests that woodland landscape females further compensate for excessive flight by redirecting resources away from somatic maintenance, resulting in reduced life spans. Our results strongly indicate that increased flight caused by changes in landscape structure may impact on key life history traits such as reproductive success and longevity.     

Enhanced immune function does not depress reproductive output

Costs of reproduction might be mediated by a physiological (resource allocation) trade-off between immune function and reproductive effort, and several recent studies have shown that an experimental increase in reproductive effort is associated with decreased immune function. Here we test the complementary prediction of this hypothesis: that increased immune function (specific antibody production) depresses reproductive output. Female European starlings (Sturnus vulgaris) were injected with a non-pathogenic antigen (sheep red blood cells) following completion of laying of their first clutch, to stimulate an in vivo humoral immune response (primary antibody production). We induced laying of a second clutch by removing the first clutch, and assessed changes in reproductive performance in individual females pre- and post-treatment. Injection of sheep red blood cells produced a significant antibody response in 96% (n=29) of treated females, with titres comparable to previous studies (range 1 to 7). However, increased antibody production did not decrease primary or secondary female reproductive effort (re-laying interval, egg size, clutch size, chick growth or fledging success), compared with control, saline-injected birds (n=22). These data do not support a simple resource allocation model for the cost of reproduction, based on a reciprocal, negative relationship between resources allocated to immune function and reproduction.     

A lengthy solution to the optimal propagule size problem in the large-bodied South American freshwater turtle, <Emphasis Type="Italic">Podocnemis unifilis</Emphasis>

In oviparous vertebrates lacking parental care, resource allocation during reproduction is a major maternal effect that may enhance female fitness. In general, resource allocation strategies are expected to follow optimality models to solve the energy trade-offs between egg size and number. Such models predict that natural selection should optimize egg size while egg number is expected to vary with female size, thus maximizing offspring fitness and consequently, maternal fitness. Deviations from optimality predictions are commonly attributed to morphological constraints imposed by female size, such as reported for small-bodied turtle species. However, whether such anatomical constraints exist in smaller-bodied females within large-bodied clades remains unstudied. Here we tested whether resource allocation of the river turtle Podocnemis unifilis (a relatively smaller member of the large-bodied Podocnemididae) follows optimality theory, and found a pattern of egg elongation in smaller females that provides evidence of morphological constraints and of a reproductive trade-off with clutch size, whereas egg width supports the existence of an optimal egg size and no trade-off. Moreover, larger females laid larger clutches composed of rounder eggs, while smaller females laid fewer and relatively more elongated eggs. Elongated eggs from smaller females have larger volume relative to female size and to round eggs of equal width. We propose that elongated eggs represent a solution to a potential morphological constraint suffered by small females. Our results suggest that larger females may optimize fitness by increasing the number of eggs, while smaller females do so by producing larger eggs. Our data supports the notion that morphological constraints are likely more widespread than previously anticipated, such that they may not be exclusive of small-bodied lineages but may also exist in large-bodied lineages.     

氯化高汞对大型溞的慢性毒性

在换水静置条件下(25℃),氯化高汞(以Hg++计)对大型溞的48小时LC50值及其可信限为13.5(±2.1)微克/升。试验个体饲以斜生栅藻单个培养,在汞含量为1—21微克/升浓度下,产仔总数经方差分析表明组间差异显著(FF0.05)。各浓度组(X)与产仔总数(Y)的关系为:Y=-51.823X+4320.960(r=-0.904,p0.01)。净增殖率(R0)随浓度升高而逐渐下降。存活和生长用方差分析及D值检验表明对照组与各浓度组之间(差异数分别小于各自的D值20.89和0.1749),没有显著差异。各浓度组对内禀增长能力(rm)和世代平均周期(T)影响不大。根据大型溞的生物学基本参数,其体长的生长模型为: lt=4.57-3.5177e-0.00838t.       

An autonomic approach for resource provisioning of cloud services

Recently, there has been a significant increase in the use of cloud-based services that are offered in software as a service (SaaS) models by SaaS providers, and irregular access of different users to these cloud services leads to fluctuations in the demand workload. It is difficult to determine the suitable amount of resources required to run cloud services in response to the varying workloads, and this may lead to undesirable states of over-provisioning and under-provisioning. In this paper, we address improvements to resource provisioning for cloud services by proposing an autonomic resource provisioning approach that is based on the concept of the control monitor-analyze-plan-execute (MAPE) loop, and we design a resource provisioning framework for cloud environments. The experimental results show that the proposed approach reduces the total cost by up to 35 %, the number of service level agreement (SLA) violations by up to 40 %, and increases the resource utilization by up to 25 % compared with the other approaches.     

Phenotypic plasticity of life-history traits in clonal and sexual fish (Poeciliopsis) at high and low densities

Models of resource allocation strategies predict an array of life-history responses of individuals living in resource-stressed versus non-stressed environments. I tested a number of these predictions using three fish strains (a sexual and two clonal strains) in high and low density treatments. To examine the plasticity of life-history traits in females raised in these two environments, I measured survival, growth, egg production, egg size, and proportion mature at 10 weeks of age. Survival was not affected by density treatment. However, both growth and overall egg production were lower in females from the high density treatments, and reproductive maturity was significantly delayed at the high density for all strains. Egg production per unit size was not affected by density in any strain, signifying that differences in the numbers of eggs produced was merely a reflection of the differences in size of fish in the two density treatments. Egg size was also unaffected by density in all strains. These results are related to models of resource allocation in stressful environments. There was a consistent pattern of increased reproductive investment in the sexual strain relative to the two clonal strains. The sexual strain matured earlier, produced more eggs per unit body weight, and had larger eggs than either clone at both densities. These results are interpreted by considering the predicted adaptive responses of these three strains to the long-term environmental differences in their natural habitats.     

Plant reproductive allocation predicts herbivore dynamics across spatial and temporal scales

Life-history theory suggests that iteroparous plants should be flexible in their allocation of resources toward growth and reproduction. Such plasticity could have consequences for herbivores that prefer or specialize on vegetative versus reproductive structures. To test this prediction, we studied the response of the cactus bug (Narnia pallidicornis) to meristem allocation by tree cholla cactus (Opuntia imbricata). We evaluated the explanatory power of demographic models that incorporated variation in cactus relative reproductive effort (RRE; the proportion of meristems allocated toward reproduction). Field data provided strong support for a single model that defined herbivore fecundity as a time-varying, increasing function of host RRE. High-RRE plants were predicted to support larger insect populations, and this effect was strongest late in the season. Independent field data provided strong support for these qualitative predictions and suggested that plant allocation effects extend across temporal and spatial scales. Specifically, late-season insect abundance was positively associated with interannual changes in cactus RRE over 3 years. Spatial variation in insect abundance was correlated with variation in RRE among five cactus populations across New Mexico. We conclude that plant allocation can be a critical component of resource quality for insect herbivores and, thus, an important mechanism underlying variation in herbivore abundance across time and space.     

Maternal effects and maternal selection arising from variation in allocation of free amino acid to eggs

Maternal provisioning can have profound effects on offspring phenotypes, or maternal effects, especially early in life. One ubiquitous form of provisioning is in the makeup of egg. However, only a few studies examine the role of specific egg constituents in maternal effects, especially as they relate to maternal selection (a standardized selection gradient reflecting the covariance between maternal traits and offspring fitness). Here, we report on the evolutionary consequences of differences in maternal acquisition and allocation of amino acids to eggs. We manipulated acquisition by varying maternal diet (milkweed or sunflower) in the large milkweed bug, Oncopeltus fasciatus. Variation in allocation was detected by examining two source populations with different evolutionary histories and life‐history response to sunflower as food. We measured amino acids composition in eggs in this 2 × 2 design and found significant effects of source population and maternal diet on egg and nymph mass and of source population, maternal diet, and their interaction on amino acid composition of eggs. We measured significant linear and quadratic maternal selection on offspring mass associated with variation in amino acid allocation. Visualizing the performance surface along the major axes of nonlinear selection and plotting the mean amino acid profile of eggs from each treatment onto the surface revealed a saddle‐shaped fitness surface. While maternal selection appears to have influenced how females allocate amino acids, this maternal effect did not evolve equally in the two populations. Furthermore, none of the population means coincided with peak performance. Thus, we found that the composition of free amino acids in eggs was due to variation in both acquisition and allocation, which had significant fitness effects and created selection. However, although there can be an evolutionary response to novel food resources, females may be constrained from reaching phenotypic optima with regard to allocation of free amino acids.