DOES TOTAL REPRODUCTIVE EFFORT EVOLVE INDEPENDENTLY OF OFFSPRING SIZE?

In all species, patterns of reproductive allocation have important fitness consequences and therefore important implications for life-history evolution. Nearly universally, theory in this field has modeled as independent the evolution of total allocation to offspring and the subsequent division of this allocation into many small versus few large offspring. Yet, some theory and a very small amount of experimental evidence suggest that these life-history traits may be evolutionarily linked. Using comparative analyses of copepod life histories, we illustrate that rather than being evolutionarily independent these traits can be linked, in this case, across a very large clade of invertebrates. Our results indicate that a more complete understanding of the evolution of these traits will require greater consideration of simultaneous allocation decisions, rather than sequential ones, and other genetic and selective mechanisms.     

Reproductive efficiency and shade avoidance plasticity under simulated competition

Plant strategy and life‐history theories make different predictions about reproductive efficiency under competition. While strategy theory suggests under intense competition iteroparous perennial plants delay reproduction and semelparous annuals reproduce quickly, life‐history theory predicts both annual and perennial plants increase resource allocation to reproduction under intense competition. We tested (1) how simulated competition influences reproductive efficiency and competitive ability (CA) of different plant life histories and growth forms; (2) whether life history or growth form is associated with CA; (3) whether shade avoidance plasticity is connected to reproductive efficiency under simulated competition. We examined plastic responses of 11 herbaceous species representing different life histories and growth forms to simulated competition (spectral shade). We found that both annual and perennial plants invested more to reproduction under simulated competition in accordance with life‐history theory predictions. There was no significant difference between competitive abilities of different life histories, but across growth forms, erect species expressed greater CA (in terms of leaf number) than other growth forms. We also found that shade avoidance plasticity can increase the reproductive efficiency by capitalizing on the early life resource acquisition and conversion of these resources into reproduction. Therefore, we suggest that a reassessment of the interpretation of shade avoidance plasticity is necessary by revealing its role in reproduction, not only in competition of plants.     

Annual plant life histories and the paradigm of resource allocation

Summary Much of life history theory follows from the idea that natural selection acts on the allocation of resources to competing and independent demographic functions. This paradigm has stimulated much research on the life histories of annual plants. Models of whole-plant resource budgets that use optimal control theory predict periods of 100% vegetative and 100% reproductive growth, sometimes with periods of mixed growth. I show here that this prediction follows from the assumption of independence of the competing vegetative and reproductive compartments. The prediction is qualitatively unchanged even after relaxing important simplifying assumptions used in most models. Although it follows naturally from the assumptions of the models, this kind of allocation pattern is unlikely to occur in many plants, because it requires that (1) leaf and flower buds can never simultaneously be carbon sinks; and (2) organs that accompany flowers, such as internodes and bracts, can never be net sources of photosynthate. Thus while resources are doubtless important for annual plants, an exclusively resource-based perspective may be inadequate to understand the evolution of their life histories. Progress in research may require models that incorporate, or are at least phenomenologically consistent with, the basic developmental reles of angiosperms.     

Life‐history trade‐offs promote the evolution of dioecy

Most dioecious plants are perennial and subject to trade‐offs between sexual reproduction and vegetative performance. However, these broader life‐history trade‐offs have not usually been incorporated into theoretical analyses of the evolution of separate sexes. One such analysis has indicated that hermaphroditism is favoured over unisexuality when female and male sex functions involve the allocation of nonoverlapping types of resources to each sex function (e.g. allocations of carbon to female function vs. allocations of nitrogen to male function). However, some dioecious plants appear to conform to this pattern of resource allocation, with different resource types allocated to female vs. male sex functions. Using an evolutionarily stable strategy approach, we show that life‐history trade‐offs between sexual reproduction and vegetative performance enable the evolution of unisexual phenotypes even when there are no direct resource‐based trade‐offs between female and male sex functions. This result might help explain the preponderance of perennial life histories among dioecious plants and why many dioecious plants with annual life histories have indeterminate growth with ongoing trade‐offs between sexual reproduction and vegetative growth.     

Life in 3-D: life history strategies in tunas, mackerels and bonitos

The scombrids (tunas, bonitos, Spanish mackerels and mackerels) sustain some of the most important fisheries in the world and their sustainable management depends on better understanding of their life history strategies. Here, we first assemble life history information on maximum size, growth, longevity, maturity, fecundity and spawning duration and interval for all scombrid species. Second we characterize their life history patterns and trait co-variation and evaluate how many principal axes of trait variation underlie scombrid life history strategies. Most of their life history variation can be explained along three axes or dimensions: size, speed, and reproductive schedule. Body size governs the first axis ranking species along a small-large continuum. The second axis was mostly influenced by time-related traits, such as longevity, growth rates, spawning duration, time between spawning events, ranking species along a slow-fast continuum of life histories. Scombrid species with the slowest life histories such as Atlantic bluefin tuna Thunnus thynnus and Atlantic mackerel Scomber scombrus tend to inhabit more temperate waters while species with faster life histories such as yellowfin tuna Thunnus albacares and short mackerel Rastrelliger brachysoma are typically found in more tropical waters. The third axis comprises the negative relationship between number of eggs produced at length of maturity and rate in gain of fecundity with size describing the schedule of reproductive allocation which reflects a fundamental trade-off between reproduction and growth. Finally, in addition we show that the life history strategies of scombrids conform more closely to the Periodic and Opportunistic strategists within the triangular model of fish life histories.     

Heuristic optimization of the general life history problem: A novel approach

Summary The general life history problem concerns the optimal allocation of resources to growth, survival and reproduction. We analysed this problem for a perennial model organism that decides once each year to switch from growth to reproduction. As a fitness measure we used the Malthusian parameterr, which we calculated from the Euler-Lotka equation. Trade-offs were incorporated by assuming that fecundity is size dependent, so that increased fecundity could only be gained by devoting more time to growth and less time to reproduction. To calculate numerically the optimalr for different growth dynamics and mortality regimes, we used a simplified version of the simulated annealing method. The major differences among optimal life histories resulted from different accumulation patterns of intrinsic mortalities resulting from reproductive costs. If these mortalities were accumulated throughout life, i.e. if they were senescent, a bangbang strategy was optimal, in which there was a single switch from growth to reproduction: after the age at maturity all resources were allocated to reproduction. If reproductive costs did not carry over from year to year, i.e. if they were not senescent, the optimal resource allocation resulted in a graded switch strategy and growth became indeterminate. Our numerical approach brings two major advantages for solving optimization problems in life history theory. First, its implementation is very simple, even for complex models that are analytically intractable. Such intractability emerged in our model when we introduced reproductive costs representing an intrinsic mortality. Second, it is not a backward algorithm. This means that lifespan does not have to be fixed at the begining of the computation. Instead, lifespan itself is a trait that can evolve. We suggest that heuristic algorithms are good tools for solving complex optimality problems in life history theory, in particular questions concerning the evolution of lifespan and senescence.     

Size and temperature in the evolution of fish life histories

Body size and temperature are the two most important variablesaffecting nearly all biological rates and times, especiallyindividual growth or production rates. By favoring an optimalmaturation age and reproductive allocation, natural selectionlinks individual growth to the mortality schedule. A recentmodel for evolution of life histories for species with indeterminategrowth, which includes most fish, successfully predicts thenumeric values of two key dimensionless numbers and the allometryof the average reproductive allocation versus maturation sizeacross species. Here we use this new model to predict the relationshipsof age-at-maturity, adult mortality and reproductive effortto environmental temperature and maturation size across species.Age-at-maturity, adult mortality and the proportion of the bodymass given to reproduction per year are predicted to show ±0.25power allometries with mass at maturity, and an exponential(Boltzmann) temperature dependence. Temperature is assumed toaffect only body size growth, so the temperature linkages ofmaturation, mortality and reproductive effort are indirect vialife history optimization; this is briefly contrasted with theidea that (for example) temperature directly affects mortality.     

Connecting proximate mechanisms and evolutionary patterns: pituitary gland size and mammalian life history

At the proximate level, hormones are known to play a critical role in influencing the life history of mammals, including humans. The pituitary gland is directly responsible for producing several hormones, including those related to growth and reproduction. Although we have a basic understanding of how hormones affect life history characteristics, we still have little knowledge of this relationship in an evolutionary context. We used data from 129 mammal species representing 14 orders to investigate the relationship between pituitary gland size and life history variation. Because pituitary gland size should be related to hormone production and action, we predicted that species with relatively large pituitaries should be associated with fast life histories, especially increased foetal and post‐natal growth rates. Phylogenetic analyses revealed that total pituitary size and the size of the anterior lobe of the pituitary significantly predicted a life history axis that was correlated with several traits including body mass, and foetal and post‐natal growth rates. Additional models directly examining the association between relative pituitary size and growth rates produced concordant results. We also found that relative pituitary size variation across mammals was best explained by an Ornstein–Uhlenbeck model of evolution, suggesting an important role of stabilizing selection. Our results support the idea that the size of the pituitary is linked to life history variation through evolutionary time. This pattern is likely due to mediating hormone levels but additional work is needed. We suggest that future investigations incorporating endocrine gland size may be critical for understanding life history evolution.     

A search for trade-offs among life history traits inDrosophila melanogaster

Summary Are there underlying developmental and physiological properties of organisms that can be used to build a general theory of life history evolution? Much of the theoretical work on the evolution of life histories is based on the premise of negative developmental and genetic correlations among life history traits. If negative correlations do not exist as a general rule then no general theory taking them into account is possible. Negative genetic correlations among life history traits can come about by antagonistic pleiotropy. One cause of antagonistic pleiotropy is cost allocation trade-offs. Since cost allocation trade-offs are due to underlying physiological constraints they are expected to be common to closely related groups. A second form of antagonistic pleiotropy is specialization of genotypes to different niches. This type of antagonistic pleiotropy is expected to be specific to each population. We looked for trade-offs in life history traits of longevity and fecundity inDrosophila melanogaster. We used a half-sib mating design and raised the offspring at two temperatures, 19°C and 25°C. Correlations between longevity and fecundity showed some evidence of antagonistic pleiotropy at high temperature with no evidence of any trade-offs at low temperature. Correlations of early and late fecundity traits did show evidence of cost allocation trade-offs at both temperatures. Antagonistic pleiotropy was also found for cross-environmental correlations of fecundity traits. We conclude that, although life history trade-offs can not be generally assumed, they are frequently found among functionally related traits. Thus, we provide guidelines for the development of general theories of life history evolution.     

The influence of juvenile and adult environments on life-history trajectories

There is increasing evidence that the environment experienced early in life can strongly influence adult life histories. It is largely unknown, however, how past and present conditions influence suites of life-history traits regarding major life-history trade-offs. Especially in animals with indeterminate growth, we may expect that environmental conditions of juveniles and adults independently or interactively influence the life-history trade-off between growth and reproduction after maturation. Juvenile growth conditions may initiate a feedback loop determining adult allocation patterns, triggered by size-dependent mortality risk. I tested this possibility in a long-term growth experiment with mouthbrooding cichlids. Females were raised either on a high-food or low-food diet. After maturation half of them were switched to the opposite treatment, while the other half remained unchanged. Adult growth was determined by current resource availability, but key reproductive traits like reproductive rate and offspring size were only influenced by juvenile growth conditions, irrespective of the ration received as adults. Moreover, the allocation of resources to growth versus reproduction and to offspring number versus size were shaped by juvenile rather than adult ecology. These results indicate that early individual history must be considered when analysing causes of life-history variation in natural populations.     

The evolution of growth rate, resource allocation and competitive ability in seeder and resprouter tree seedlings

Many woody plant species in fire disturbed communities survive disturbance events by resprouting. The resprouting life history is predicted to be costly to plants as resources are diverted into storage for post-fire regrowth rather than allocated to current growth, and resprouting species typically grow more slowly than seeder species (species that do not resprout after disturbance events). Differences in allocation to current growth are also predicted to make resprouter species poorer competitors compared to seeder species. We tested the predictions that the evolution of a resprouter life history is associated with slow growth, increased allocation to storage, and low competitive ability in woody plant seedlings. We grew eight phylogenetically independent pairs of seeder and resprouter species in competition and no competition treatments in a field experiment near Sydney, Australia. The presence of competitors reduced plant growth rates across taxa and fire response life histories. However, relative to seeder species, resprouter species were not slower growing, they did not allocate more resources to storage, and they did not have lower competitive abilities. We propose that differences in resource allocation to storage are not responsible for differences in growth rate and competitive ability. Rather, growth rate and competitive ability in seedlings are associated with key aspects of plant life history such as life-span and body size at maturity. These traits that are sometimes, but not always, related to fire response life histories.     

Life-history evolution in guppies viii: the demographics of density regulation in guppies (poecilia reticulata)

In prior research, we found the way guppy life histories evolve in response to living in environments with a high or low risk of predation is consistent with life-history theory that assumes no density dependence. We later found that guppies from high-predation environments experience higher mortality rates than those from low-predation environments, but the increased risk was evenly distributed across all age/size classes. Life-history theory that assumes density-independent population growth predicts that life histories will not evolve under such circumstances, yet we have shown with field introduction experiments that they do evolve. However, theory that incorporates density regulation predicts this pattern of mortality can result in the patterns of life-history evolution we had observed. Here we report on density manipulation experiments performed in populations of guppies from low-predation environments to ask whether natural populations normally experience density regulation and, if so, to characterize the short-term demographic changes that underlie density regulation. Our experiments reveal that these populations are density regulated. Decreased density resulted in higher juvenile growth, decreased juvenile mortality rates, and increased reproductive investment by adult females. Increased density causes reduced offspring size, decreased fat storage by adult females, and increased adult mortality.     

Evolution of resource allocation between growth and reproduction in animals with indeterminate growth

We review the recent theoretical developments explaining the evolution of age-schedules of reproduction in animals with indeterminate growth. Indeterminate growth, i.e. growth that continues past maturation and may continue until the end of life, is characteristic for a large number of invertebrate taxa (e.g. clams, cladocerans and crayfish) and ‘lower’ vertebrate taxa (e.g. fish, amphibians, lizards and snakes). Many plants also exhibit indeterminate growth, and we liberally include studies focused on plants when they can be interpreted in terms of animal life histories. We focus on different measures used to determine the fittest life histories, on indeterminate growth as a problem of resource allocation and on the effects of environment to the evolution of the resource allocation schemes.     

The impact of parasitism on resource allocation in a fungal host: the case of Cryphonectria parasitica and its mycovirus,Cryphonectria Hypovirus 1

Parasites are known to profoundly affect resource allocation in their host. In order to investigate the effects of Cryphonectria Hypovirus 1 (CHV1) on the life‐history traits of its fungal host Cryphonectria parasitica, an infection matrix was completed with the cross‐infection of six fungal isolates by six different viruses. Mycelial growth, asexual sporulation, and spore size were measured in the 36 combinations, for which horizontal and vertical transmission of the viruses was also assessed. As expected by life‐history theory, a significant negative correlation was found between host somatic growth and asexual reproduction in virus‐free isolates. Interestingly this trade‐off was found to be positive in infected isolates, illustrating the profound changes in host resource allocation induced by CHV1 infection. A significant and positive relationship was also found in infected isolates between vertical transmission and somatic growth. This last relationship suggests that in this system, high levels of virulence could be detrimental to the vertical transmission of the parasite. Those results underscore the interest of studying host–parasite interaction within the life‐history theory framework, which might permit a more accurate understanding of the nature of the modifications triggered by parasite infection on host biology.     

Quantifying multivariate plasticity: genetic variation in resource acquisition drives plasticity in resource allocation to components of life history

Acquisition and allocation of resources are central to life‐history theory. However, empirical work typically focuses only on allocation despite the fact that relationships between fitness components may be governed by differences in the ability of individuals to acquire resources across environments. Here, we outline a statistical framework to partition the genetic basis of multivariate plasticity into independent axes of genetic variation, and quantify for the first time, the extent to which specific traits drive multitrait genotype–environment interactions. Our framework generalises to analyses of plasticity, growth and ageing. We apply this approach to a unique, large‐scale, multivariate study of acquisition, allocation and plasticity in the life history of the cricket, Gryllus firmus. We demonstrate that resource acquisition and allocation are genetically correlated, and that plasticity in trade‐offs between allocation to components of fitness is 90% dependent on genetic variance for total resource acquisition. These results suggest that genotype–environment effects for resource acquisition can maintain variation in life‐history components that are typically observed in the wild.     

Evolution of alternative insect life histories in stochastic seasonal environments

Deterministic seasonality can explain the evolution of alternative life history phenotypes (i.e., life history polyphenism) expressed in different generations emerging within the same year. However, the influence of stochastic variation on the expression of such life history polyphenisms in seasonal environments is insufficiently understood. Here, we use insects as a model and explore (1) the effects of stochastic variation in seasonality and (2) the life cycle on the degree of life history differentiation among the alternative developmental pathways of direct development and diapause (overwintering), and (3) the evolution of phenology. With numerical simulation, we determine the values of development (growth) time, growth rate, body size, reproductive effort, adult life span, and fecundity in both the overwintering and directly developing generations that maximize geometric mean fitness. The results suggest that natural selection favors the expression of alternative life histories in the alternative developmental pathways even when there is stochastic variation in seasonality, but that trait differentiation is affected by the developmental stage that overwinters. Increasing environmental unpredictability induced a switch to a bet‐hedging type of life history strategy, which is consistent with general life history theory. Bet‐hedging appeared in our study system as reduced expression of the direct development phenotype, with associated changes in life history phenotypes, because the fitness value of direct development is highly variable in uncertain environments. Our main result is that seasonality itself is a key factor promoting the evolution of seasonally polyphenic life histories but that environmental stochasticity may modulate the expression of life history phenotypes.     

The importance of growth and mortality costs in the evolution of the optimal life history

A central assumption of life history theory is that the evolution of the component traits is determined in part by trade-offs between these traits. Whereas the existence of such trade-offs has been well demonstrated, the relative importance of these remains unclear. In this paper we use optimality theory to test the hypothesis that the trade-off between present and future fecundity induced by the costs of continued growth is a sufficient explanation for the optimal age at first reproduction, alpha, and the optimal allocation to reproduction, G, in 38 populations of perch and Arctic char. This hypothesis is rejected for both traits and we conclude that this trade-off, by itself, is an insufficient explanation for the observed values of alpha and G. Similarly, a fitness function that assumes a mortality cost to reproduction but no growth cost cannot account for the observed values of alpha. In contrast, under the assumption that fitness is maximized, the observed life histories can be accounted for by the joint action of trade-offs between growth and reproductive allocation and between mortality and reproductive allocation (Individual Juvenile Mortality model). Although the ability of the growth/mortality model to fit the data does not prove that this is the mechanism driving the evolution of the optimal age at first reproduction and allocation to reproduction, the fit does demonstrate that the hypothesis is consistent with the data and hence cannot at this time be rejected. We also examine two simpler versions of this model, one in which adult mortality is a constant proportion of juvenile mortality [Proportional Juvenile Mortality (PJM) model] and one in which the proportionality is constant within but not necessarily between species [Specific Juvenile Mortality (SSJM) model]. We find that the PJM model is unacceptable but that the SSJM model produces fits suggesting that, within the two species studied, juvenile mortality is proportional to adult mortality but the value differs between the two species.     

Twelve fundamental life histories evolving through allocation‐dependent fecundity and survival

An organism's life history is closely interlinked with its allocation of energy between growth and reproduction at different life stages. Theoretical models have established that diminishing returns from reproductive investment promote strategies with simultaneous investment into growth and reproduction (indeterminate growth) over strategies with distinct phases of growth and reproduction (determinate growth). We extend this traditional, binary classification by showing that allocation‐dependent fecundity and mortality rates allow for a large diversity of optimal allocation schedules. By analyzing a model of organisms that allocate energy between growth and reproduction, we find twelve types of optimal allocation schedules, differing qualitatively in how reproductive allocation increases with body mass. These twelve optimal allocation schedules include types with different combinations of continuous and discontinuous increase in reproduction allocation, in which phases of continuous increase can be decelerating or accelerating. We furthermore investigate how this variation influences growth curves and the expected maximum life span and body size. Our study thus reveals new links between eco‐physiological constraints and life‐history evolution and underscores how allocation‐dependent fitness components may underlie biological diversity.     

Life history trade-offs are more pronounced for a noninvasive,native butterfly compared to its invasive,exotic congener

Life history trade-offs are ubiquitous in nature. Life history theory posits that these trade-offs arise from individuals having limited resources to allocate toward all vital functions, such as survival, growth and reproduction. These trade-offs position most species along a slow-fast life history continuum, where individuals with slow life histories often have higher survival at the cost of delayed reproduction and individuals with fast life histories often live faster and die younger. However, these trade-offs are sometimes less obvious for invasive species. Here, we constructed age-based population models to compare life history strategies and trade-offs between the noninvasive, native mustard white and invasive, exotic cabbage white (Pieris spp.) butterflies. We found that the cabbage white has faster larval growth and higher fecundity at younger ages, suggesting it has a fast life history compared to the mustard white. However, cabbage white also has higher adult survival at younger ages, suggesting that it experiences weaker trade-offs among vital rates than its native counterpart. Our study illustrates the importance of demographic studies in evaluating life history strategies among congener species with different population histories, and emphasizes the many advantages experienced by invasive species in their novel environments.     

Evolution of Rotifer Life Histories

When compared to most other multicellular animals, rotifers are all relatively small, short-lived and fast-reproducing organisms. However among and within different rotifer species there is a large variation in life history patterns. This review accounts for such variation in rotifers, with a strong focus on monogonont rotifers. As the life cycle of monogonont rotifers involves both asexual and sexual reproduction, life history patterns can be examined on the level of the genetic individual, which includes all asexual females, sexual females and males that originated from one resting egg. This concept has been applied successfully in many areas, for example in predicting optimal levels of mictic reproduction or sex allocation theory. The benefits and implications of the view of the genetic individual are discussed in detail. Rotifer life histories can also be viewed on the level of physiological individuals. A large part of this review deals with the life histories of individual amictic females and addresses life history traits like body size, egg size and resource allocation patterns. It asks which trade-offs exist among those traits, how these traits change under the influence of environmental factors like food availability or temperature, and whether these changes can be interpreted as adaptive.