Senescence and the stress axis: a constraint or a trade-off of reproduction in mammals with fast and slow life histories.

A critical time in the life of a male occurs at reproduction, when his behaviour, physiology and resources must be brought to bear for the central purpose of his life-propagating his genes. We ask whether reproduction results in dysfunction of the stress axis, is linked to life history, and causes senescence. We assessed if deterioration in the axis underlies variation in reproductive lifespan in males of five species of North American ground squirrels whose life history varies from near semelparity to iteroparity. The most stressful and energy-demanding time occurs in spring during the intense 2-3 week breeding competition just after arousal from hibernation. We compared their stress axis functioning before and after the mating period using a hormonal challenge protocol. We found no evidence of stress axis dysfunction after reproduction in any species nor was there a relationship between reproductive lifespan and stress axis functional deterioration. Moreover, there was no consistent relationship between free cortisol levels and downstream measures (energy mobilization, haematology, immunity and body indices of condition). Thus, stress axis function was not traded off to promote reproduction irrespective of life history and lifespan, and we conclude that it is a prerequisite for life. Hence, it functions as a constraint and does not undergo senescence.

Feeding after spawning and energy balance at spawning are associated with repeat spawning interval in steelhead trout.

Consecutive and skip repeat spawning (1- or ≥2-year spawning interval) life histories commonly occur in seasonally breeding iteroparous fishes. Spawning interval variation is driven by energetic status and impacts fisheries management. In salmonids, energetic status (either absolute level of energy reserves or the rate of change of energy reserves, i.e., energy balance) is thought to determine reproductive trajectory during a critical period ∼1 year prior to initial spawning. However, information on repeat spawners is lacking. To examine the timing and the aspects of energetic status that regulate repeat spawning interval, female steelhead trout (Oncorhynchus mykiss) were fasted for 10 weeks after spawning and then fed ad libitum and compared to ad libitum fed controls. Plasma growth hormone (GH) and insulin-like growth factor-I (IGF-I) levels were measured to assess long-term energy balance. Plasma estradiol levels showed that some fish in both groups initiated a consecutive spawning cycle. In fasted fish, GH was lower at spawning in consecutive versus skip spawners. In consecutive spawners, GH was higher at spawning in fed versus fasted fish. These results suggest that fish with a less negative energy balance at spawning initiated reproductive development in the absence of feeding, but that feeding during the post-spawning period enabled initiation of reproduction in some fish with a more negative energy balance at spawning. Thus, both energy balance at spawning and feeding after spawning regulated reproductive schedules. These results show that the critical period model of salmonid maturation applies to regulation of repeat spawning, and that the reproductive decision window extends into the first 10 weeks after spawning.

Effects of post-spawning ration restriction on reproductive development and the growth hormone/insulin-like growth factor-1 axis in female rainbow trout (Oncorhynchus mykiss).

In iteroparous female salmonids, the growth and reproductive endocrine axes interact during the period after spawning. Energy depletion due to pre-spawn fasting, migration, and ovarian development must be restored, and the next reproductive cycle is initiated in consecutively maturing fish. In the natural environment, food availability is often limited during the post-spawn period. To investigate the growth and reproductive endocrinology of the post-spawn period, we sampled female rainbow trout over the 30 weeks following their first spawning. Fish were fasted for 2 months prior to spawning, then fed a standard or a restricted ration. Analysis was confined to reproductive fish. Plasma estradiol-17ß decreased during the 8 weeks following spawning and then began increasing in both ration groups and was lower in feed-restricted versus standard ration fish from 8 weeks onward. Plasma insulin-like growth factor-1 increased over the same period and then remained constant in both ration groups and was lower in feed-restricted versus standard ration fish from week 8 to week 30. Plasma growth hormone decreased following spawning in standard ration fish and became elevated in feed-restricted versus standard ration fish at 20- and 30-weeks post-spawn. Growth rates, condition factor, and muscle lipid levels were higher in standard ration versus feed-restricted fish within 2-4 weeks after spawning. These results suggest that two phases occurred during the post-spawn period: recovery from spawning and restoration of energy reserves over weeks 0 to 8, followed by adjustment of the growth and reproductive endocrine axes to ration level over weeks 8 to 30.

Life history variation is maintained by fitness trade-offs and negative frequency-dependent selection.

The maintenance of diverse life history strategies within and among species remains a fundamental question in ecology and evolutionary biology. By using a near-complete 16-year pedigree of 12,579 winter-run steelhead (Oncorhynchus mykiss) from the Hood River, Oregon, we examined the continued maintenance of two life history traits: the number of lifetime spawning events (semelparous vs. iteroparous) and age at first spawning (2-5 years). We found that repeat-spawning fish had more than 2.5 times the lifetime reproductive success of single-spawning fish. However, first-time repeat-spawning fish had significantly lower reproductive success than single-spawning fish of the same age, suggesting that repeat-spawning fish forego early reproduction to devote additional energy to continued survival. For single-spawning fish, we also found evidence for a fitness trade-off for age at spawning: older, larger males had higher reproductive success than younger, smaller males. For females, in contrast, we found that 3-year-old fish had the highest mean lifetime reproductive success despite the observation that 4- and 5-year-old fish were both longer and heavier. This phenomenon was explained by negative frequency-dependent selection: as 4- and 5-year-old fish decreased in frequency on the spawning grounds, their lifetime reproductive success became greater than that of the 3-year-old fish. Using a combination of mathematical and individual-based models parameterized with our empirical estimates, we demonstrate that both fitness trade-offs and negative frequency-dependent selection observed in the empirical data can theoretically maintain the diverse life history strategies found in this population.

Trade-off between seed dispersal in space and time.

Seed movement and delayed germination have long been thought to represent alternative risk-spreading strategies, but current evidence covers limited scales and yields mixed results. Here we present the first global-scale test of a negative correlation between dispersal and dormancy. The result demonstrates a strong and consistent pattern that species with dormant seeds have reduced spatial dispersal, also in the context of life-history traits such as seed mass and plant lifespan. Long-lived species are more likely to have large, non-dormant seeds that are dispersed far. Our findings provide robust support for the theoretical prediction of a dispersal trade-off between space and time, implying that a joint consideration of risk-spreading strategies is imperative in studying plant life-history evolution. The bet-hedging patterns in the dispersal-dormancy correlation and the associated reproductive traits have implications for biodiversity conservation, via prediction of which plant groups would be most impacted in the changing era.

Life history variation in an invasive plant is associated with climate and recent colonization of a specialist herbivore.

PREMISE: Spatial variation in selective pressures can lead to intraspecific variation in life history, favoring some life histories and constraining others depending on the vulnerability of life stages. We examined how spatial variation in herbivory and climate influences flowering size and the occurrence of semelparity (reproducing once) versus iteroparity (reproducing multiple times) in the introduced range of an invasive plant, houndstongue (Cynoglossum officinale). Houndstongue is a short-lived, semelparous perennial in its native range. In its introduced range, we previously documented increased rates of iteroparity and a higher median threshold flowering size compared to the native range. We hypothesized that the recent introduction of a specialist biocontrol insect (a root-boring weevil, Mogulones crucifer) would decrease threshold flowering size and reduce the proportion of iteroparous plants because M. crucifer preferentially attacks large individuals and may reduce overwinter survival. METHODS: We surveyed 24 sites across the northwestern United States to quantify the frequency of semelparity versus iteroparity and to estimate weevil abundance and used demographic data collected from six sites to estimate median threshold flowering size. RESULTS: We found that sites with greater winter precipitation and no weevils had a greater proportion of iteroparous plants. Sites with higher weevil attack had a lower median threshold flowering size. CONCLUSIONS: The variation in frequency of flowering and threshold flowering size that we documented in North American houndstongue populations, and the relationships between this variation and herbivory and climate, provide evidence for how selective pressures covary with the life histories of invasive plants.

Co-inheritance of sea age at maturity and iteroparity in the Atlantic salmon vgll3 genomic region.

Co-inheritance in life-history traits may result in unpredictable evolutionary trajectories if not accounted for in life-history models. Iteroparity (the reproductive strategy of reproducing more than once) in Atlantic salmon (Salmo salar) is a fitness trait with substantial variation within and among populations. In the Teno River in northern Europe, iteroparous individuals constitute an important component of many populations and have experienced a sharp increase in abundance in the last 20 years, partly overlapping with a general decrease in age structure. The physiological basis of iteroparity bears similarities to that of age at first maturity, another life-history trait with substantial fitness effects in salmon. Sea age at maturity in Atlantic salmon is controlled by a major locus around the vgll3 gene, and we used this opportunity demonstrate that these two traits are co-inherited around this genome region. The odds ratio of survival until second reproduction was up to 2.4 (1.8-3.5 90% CI) times higher for fish with the early-maturing vgll3 genotype (EE) compared to fish with the late-maturing genotype (LL). The L allele was dominant in individuals remaining only one year at sea before maturation, but the dominance was reversed, with the E allele being dominant in individuals maturing after two or more years at sea. Post hoc analysis indicated that iteroparous fish with the EE genotype had accelerated growth prior to first reproduction compared to first-time spawners, across all age groups, whereas this effect was not detected in fish with the LL genotype. These results broaden the functional link around the vgll3 genome region and help us understand constraints in the evolution of life-history variation in salmon. Our results further highlight the need to account for genetic correlations between fitness traits when predicting demographic changes in changing environments.

Fast-slow continuum and reproductive strategies structure plant life-history variation worldwide.

The identification of patterns in life-history strategies across the tree of life is essential to our prediction of population persistence, extinction, and diversification. Plants exhibit a wide range of patterns of longevity, growth, and reproduction, but the general determinants of this enormous variation in life history are poorly understood. We use demographic data from 418 plant species in the wild, from annual herbs to supercentennial trees, to examine how growth form, habitat, and phylogenetic relationships structure plant life histories and to develop a framework to predict population performance. We show that 55% of the variation in plant life-history strategies is adequately characterized using two independent axes: the fast-slow continuum, including fast-growing, short-lived plant species at one end and slow-growing, long-lived species at the other, and a reproductive strategy axis, with highly reproductive, iteroparous species at one extreme and poorly reproductive, semelparous plants with frequent shrinkage at the other. Our findings remain consistent across major habitats and are minimally affected by plant growth form and phylogenetic ancestry, suggesting that the relative independence of the fast-slow and reproduction strategy axes is general in the plant kingdom. Our findings have similarities with how life-history strategies are structured in mammals, birds, and reptiles. The position of plant species populations in the 2D space produced by both axes predicts their rate of recovery from disturbances and population growth rate. This life-history framework may complement trait-based frameworks on leaf and wood economics; together these frameworks may allow prediction of responses of plants to anthropogenic disturbances and changing environments.

Prey productivity and predictability drive different axes of life-history variation in carnivorous marsupials.

Variation in life-history strategies has usually been characterized as a single fast-slow continuum of life-history variation, in which mean lifespan increases with age at maturity as reproductive output at each breeding event declines. Analyses of plants and animals suggest that strategies of reproductive timing can vary on an independent axis, with iteroparous species at one extreme and semelparous species at the other. Insectivorous marsupials in the Family Dasyuridae have an unusually wide range of life-history strategies on both purported axes. We test and confirm that reproductive output and degree of iteroparity are independent in females across species. Variation in reproductive output per episode is associated with mean annual rainfall, which predicts food availability. Position on the iteroparity-semelparity axis is not associated with annual rainfall, but species in regions of unpredictable rainfall have longer maximum lifespans, more potential reproductive events per year, and longer breeding seasons. We suggest that these two axes of life-history variation arise because reproductive output is limited by overall food availability, and selection for high offspring survival favours concentrated breeding in seasonal environments. Longer lifespans are favoured when reproductive opportunities are dispersed over longer periods in environments with less predictable food schedules.

Estimating demographic contributions to effective population size in an age-structured wild population experiencing environmental and demographic stochasticity.

A population's effective size (Ne ) is a key parameter that shapes rates of inbreeding and loss of genetic diversity, thereby influencing evolutionary processes and population viability. However, estimating Ne , and identifying key demographic mechanisms that underlie the Ne to census population size (N) ratio, remains challenging, especially for small populations with overlapping generations and substantial environmental and demographic stochasticity and hence dynamic age-structure. A sophisticated demographic method of estimating Ne /N, which uses Fisher's reproductive value to account for dynamic age-structure, has been formulated. However, this method requires detailed individual- and population-level data on sex- and age-specific reproduction and survival, and has rarely been implemented. Here, we use the reproductive value method and detailed demographic data to estimate Ne /N for a small and apparently isolated red-billed chough (Pyrrhocorax pyrrhocorax) population of high conservation concern. We additionally calculated two single-sample molecular genetic estimates of Ne to corroborate the demographic estimate and examine evidence for unobserved immigration and gene flow. The demographic estimate of Ne /N was 0.21, reflecting a high total demographic variance (σ2dg) of 0.71. Females and males made similar overall contributions to σ2dg. However, contributions varied among sex-age classes, with greater contributions from 3 year-old females than males, but greater contributions from ≥5 year-old males than females. The demographic estimate of Ne was ~30, suggesting that rates of increase of inbreeding and loss of genetic variation per generation will be relatively high. Molecular genetic estimates of Ne computed from linkage disequilibrium and approximate Bayesian computation were approximately 50 and 30, respectively, providing no evidence of substantial unobserved immigration which could bias demographic estimates of Ne . Our analyses identify key sex-age classes contributing to demographic variance and thus decreasing Ne /N in a small age-structured population inhabiting a variable environment. They thereby demonstrate how assessments of Ne can incorporate stochastic sex- and age-specific demography and elucidate key demographic processes affecting a population's evolutionary trajectory and viability. Furthermore, our analyses show that Ne for the focal chough population is critically small, implying that management to re-establish genetic connectivity may be required to ensure population viability.

Costs of reproduction can explain the correlated evolution of semelparity and egg size: theory and a test with salmon.

Species' life history traits, including maturation age, number of reproductive bouts, offspring size and number, reflect adaptations to diverse biotic and abiotic selection pressures. A striking example of divergent life histories is the evolution of either iteroparity (breeding multiple times) or semelparity (breed once and die). We analysed published data on salmonid fishes and found that semelparous species produce larger eggs, that egg size and number increase with salmonid body size among populations and species and that migratory behaviour and parity interact. We developed three hypotheses that might explain the patterns in our data and evaluated them in a stage-structured modelling framework accounting for different growth and survival scenarios. Our models predict the observation of small eggs in iteroparous species when egg size is costly to maternal survival or egg number is constrained. By exploring trait co-variation in salmonids, we generate new hypotheses for the evolution of trade-offs among life history traits.

Effective number of breeders, effective population size and their relationship with census size in an iteroparous species, Salvelinus fontinalis.

The relationship between the effective number of breeders (Nb) and the generational effective size (Ne) has rarely been examined empirically in species with overlapping generations and iteroparity. Based on a suite of 11 microsatellite markers, we examine the relationship between Nb, Ne and census population size (Nc) in 14 brook trout (Salvelinus fontinalis) populations inhabiting 12 small streams in Nova Scotia and sampled at least twice between 2009 and 2015. Unbiased estimates of Nb obtained with individuals of a single cohort, adjusted on the basis of age at first maturation (α) and adult lifespan (AL), were from 1.66 to 0.24 times the average estimates of Ne obtained with random samples of individuals of mixed ages (i.e. [Formula: see text]). In turn, these differences led to adjusted Ne estimates that were from nearly five to 0.7 times the estimates derived from mixed-aged individuals. These differences translate into the same range of variation in the ratio of effective to census population size [Formula: see text] within populations. Adopting [Formula: see text] as the more precise and unbiased estimates, we found that these brook trout populations differ markedly in their effective to census population sizes (range approx. 0.3 to approx. 0.01). Using AgeNe, we then showed that the variance in reproductive success or reproductive skew varied among populations by a factor of 40, from Vk/k ≈ 5 to 200. These results suggest wide differences in population dynamics, probably resulting from differences in productivity affecting the intensity of competition for access to mates or redds, and thus reproductive skew. Understanding the relationship between Ne, Nb and Nc, and how these relate to population dynamics and fluctuations in population size, are important for the design of robust conservation strategies in small populations with overlapping generations and iteroparity.

Dispersal, dormancy and life-history tradeoffs at the individual, population and species levels in southern African Asteraceae.

Dispersal and dormancy are important risk-reducing strategies in unpredictable environments. Negative covariation between these strategies is theoretically expected, but empirical evidence is limited and inconsistent. Moreover, covariation may be affected by other life-history traits and may vary across levels of biological organization. We assessed dispersal (vertical fall time of fruits, a proxy for wind dispersal ability) and dormancy (germination fractions measured during germination trials) in populations of 15 annual and 12 perennial wind-dispersed species in six Asteraceae genera from South Africa. Dormancy was higher in annuals than in perennials, whereas fall time was largely determined by evolutionary history. Controlling for phylogeny, dispersal and dormancy was negatively associated across species and life-history categories. Negative covariation between dispersal and dormancy was not evident at either the individual level (except for seed heteromorphic species) or the population level. Our study provides rare empirical support for the theoretical expectation of tradeoffs between dormancy and the alternative risk-reducing strategies, perenniality and dispersal, but refutes the expectation of increased dispersability in perennials. Although negative covariation between dispersal and dormancy at the species level appears not to be a simple consequence of upscaling individual-level mechanistic tradeoffs, our findings suggest that selection for one strategy may constrain evolution of the other.

Fatty-acid profiles of white muscle and liver in stream-maturing steelhead trout Oncorhynchus mykiss from early migration to kelt emigration.

The profiles of specific fatty acids (FA) in white muscle and liver of fasting steelhead trout Oncorhynchus mykiss were evaluated at three periods during their prespawning migration and at kelt emigration in the Snake-Columbia River of Washington, Oregon and Idaho, to improve the understanding of energy change. Twenty-seven FAs were identified; depletion of 10 of these was positively correlated in liver and white muscle of prespawning O. mykiss. To observe relative changes in FA content more accurately over sampling intervals, the lipid fraction of tissues was used to normalize the quantity of individual FA to an equivalent tissue wet mass. Saturated and monounsaturated FAs were depleted between upstream migration in September and kelt emigration in June, whereas polyunsaturated FAs were more conserved. Liver was depleted of FAs more rapidly than muscle. Three FAs were detected across all sampling intervals: 16:0, 18:1 and 22:6n3, which are probably structurally important to membranes. When structurally important FAs of O. mykiss are depleted to provide energy, physiological performance and survival may be affected.

Life-history diversity and its importance to population stability and persistence of a migratory fish: steelhead in two large North American watersheds.

Life-history strategies can buffer individuals and populations from environmental variability. For instance, it is possible that asynchronous dynamics among different life histories can stabilize populations through portfolio effects. Here, we examine life-history diversity and its importance to stability for an iconic migratory fish species. In particular, we examined steelhead (Oncorhynchus mykiss), an anadromous and iteroparous salmonid, in two large, relatively pristine, watersheds, the Skeena and Nass, in north-western British Columbia, Canada. We synthesized life-history information derived from scales collected from adult steelhead (N = 7227) in these watersheds across a decade. These migratory fishes expressed 36 different manifestations of the anadromous life-history strategy, with 16 different combinations of freshwater and marine ages, 7·6% of fish performing multiple spawning migrations, and up to a maximum of four spawning migrations per lifetime. Furthermore, in the Nass watershed, various life histories were differently prevalent through time - three different life histories were the most prevalent in a given year, and no life history ever represented more than 45% of the population. These asynchronous dynamics among life histories decreased the variability of numerical abundance and biomass of the aggregated population so that it was > 20% more stable than the stability of the weighted average of specific life histories: evidence of a substantial portfolio effect. Year of ocean entry was a key driver of dynamics; the median correlation coefficient of abundance of life histories that entered the ocean the same year was 2·5 times higher than the median pairwise coefficient of life histories that entered the ocean at different times. Simulations illustrated how different elements of life-history diversity contribute to stability and persistence of populations. This study provides evidence that life-history diversity can dampen fluctuations in population abundances and biomass via portfolio effects. Conserving genetic integrity and habitat diversity in these and other large watersheds can enable a diversity of life histories that increases population and biomass stability in the face of environmental variability.

Changing reproductive effort within a semelparous reproductive episode.

UNLABELLED: • PREMISE OF THE STUDY: Life-history theory predicts a trade-off between current and future reproduction for iteroparous organisms-as individuals age, the expected value of future reproduction declines, and thus reproductive effort is expected to be higher in later clutches than in earlier. In contrast, models explaining the evolution of semelparity treat semelparous reproduction as instantaneous, with no scope for intraindividual variation. However, semelparous reproduction is also extended, but over shorter time scales; whether there are similar age- or stage-specific changes in reproductive effort within a semelparous episode is unclear. In this study, we assessed whether semelparous individuals increase reproductive effort as residual reproductive value declines by comparing the reproductive phenotype of flowers at five different floral positions along a main inflorescence.• METHODS: Using the herbaceous monocarp Lobelia inflata, we conducted a longitudinal study of 409 individuals including both laboratory and field populations over three seasons. We recorded six reproductive traits-including the length of three phenological intervals as well as fruit size, seed size, and seed number-for all plants across floral positions produced throughout the reproductive episode.• KEY RESULTS: We found that while the rate of flower initiation did not change, flowers at distal (late) floral positions developed more quickly and contained larger seed than flowers at basal (early) floral positions did.• CONCLUSIONS: Our results were consistent with the hypothesis that, like iteroparous organisms, L. inflata increases reproductive effort in response to low residual reproductive value.

Simple life-history traits explain key effective population size ratios across diverse taxa.

Effective population size (Ne) controls both the rate of random genetic drift and the effectiveness of selection and migration, but it is difficult to estimate in nature. In particular, for species with overlapping generations, it is easier to estimate the effective number of breeders in one reproductive cycle (Nb) than Ne per generation. We empirically evaluated the relationship between life history and ratios of Ne, Nb and adult census size (N) using a recently developed model (agene) and published vital rates for 63 iteroparous animals and plants. Nb/Ne varied a surprising sixfold across species and, contrary to expectations, Nb was larger than Ne in over half the species. Up to two-thirds of the variance in Nb/Ne and up to half the variance in Ne/N was explained by just two life-history traits (age at maturity and adult lifespan) that have long interested both ecologists and evolutionary biologists. These results provide novel insights into, and demonstrate a close general linkage between, demographic and evolutionary processes across diverse taxa. For the first time, our results also make it possible to interpret rapidly accumulating estimates of Nb in the context of the rich body of evolutionary theory based on Ne per generation.

Metabolic endocrine factors involved in spawning recovery and rematuration of iteroparous female rainbow trout (Oncorhynchus mykiss).

To determine how energy balance affects metabolic hormones hypothesized to play a role in the onset of a new reproductive cycle in iteroparous salmonids, food availability after spawning was restricted in female rainbow trout. These fish were compared with a control group that was fed a standard brood stock ration. Bodyweight, length, and muscle lipid content were determined, and blood was collected from fish at regular intervals; a subset of fish from each group was sacrificed at each sampling time for the collection of liver and ovary tissue, and to calculate hepatosomatic index (HSI) and gonadosomatic index (GSI). Plasma hormone levels were quantified by radioimmunoassay, and tissue gene expression levels were analyzed using q-RT-PCR. The experiment was conducted twice, using two-year-old and three-year-old post-spawned fish. Food-restriction arrested ovarian growth and development within 15-20 weeks, as evidenced by lower GSI in restricted-ration fish. Food restriction also reduced Fulton's condition factor, muscle lipid content, and specific growth rate from one month onward, and reduced HSI after 3 months. In the liver, insulin-like growth factor (igf1 and igf2) gene expression was reduced in three-year-old food-restricted fish within 2 months; however, no effect of ration on igf1 or igf2 expression was detected in two-year-old fish. In both years, IGF binding protein-1 (igfbp1) gene expression decreased over time in both treatment groups. Liver leptin (slepA1) gene expression was lower in two-year-old food-restricted fish at 4 months. These results show that this feed restriction regime arrested reproductive development and affected factors associated with energy balance purported to play a role in initiating reproductive development within 2-4months after spawning.

Adaptive delayed reproduction in a 2-dimensional discrete-time competition model.

This paper studies a 2-dimensional discrete-time competition model of Ricker type with reproductive delay. The model is examined under the assumption that species 1 and 2 have the same properties except that a fraction η of species 1 individuals delays the initiation of reproduction. This assumption ensures that species 1 is dominated by species 2 in the sense that species 2 is increasing whenever species 1 is increasing. It is shown that, even under this assumption, delayed reproduction can be adaptive, i.e. species 1 can invade the monoculture system of species 2 while species 2 cannot invade the monoculture system of species 1, if the population is fluctuating. The result is obtained by analytically examining the species invasibility at boundary 2-cycles, whose coordinates can be estimated by assuming η≈0.

Modalities of aging in organisms with different strategies of resource allocation.

Although the progress of aging research relies heavily on a theoretical framework, today there is no consensus on many critical questions in aging biology. I hypothesize that a systematic analysis of the intersection of different evolutionary mechanisms of aging with diverse resource allocation strategies in different organisms may reconcile aging hypotheses. The application of disposable soma, mutation accumulation, antagonistic pleiotropy, and life-history theory is considered across organisms with asexual reproduction, organisms with sexual reproduction and indeterminate growth in different conditions of extrinsic mortality, and organisms with determinate growth, with endotherms/homeotherms as a subgroup. This review demonstrates that different aging mechanisms are complementary to each other, and in organisms with different resource allocation strategies they form aging modalities ranging from immortality to suicidal programs. It also revamps the role of growth arrest in aging. Growth arrest evolved in many different groups of organisms as a result of resource reallocation from growth to reproduction (e.g., semelparous animals, holometabolic insects), or from growth to nutrient storage (endotherms/homeotherms). Growth arrest in different animal lineages has similar molecular mechanisms and similar consequences for longevity due to the conflict between growth-promoting and growth-suppressing programs and suppression of regenerative capacity.