Trade‐offs between growth and reproduction: an analysis of the quantitative genetic evidence

The assumption of a trade‐off between development time and fecundity, resulting from a positive correlation between body size and fecundity and between body size and development time, is a common feature of life history models. The present paper examines the evidence for such a trade‐off as indicated by genetic correlations between traits. The genetic covariances between traits are derived using a model in which maturation occurs when the organism achieves a genetically variable size threshold, and fecundity is an allometric function of body size with one genetically variable parameter (excluding body size itself). This model predicts that the heritabilities of the life history traits (growth rate, development time, fecundity) will not necessarily be less than the heritability of adult size (i.e. morphological traits). It is shown that if growth rate is genetically correlated with adult size then it is not possible, in general, to predict the sign of the genetic correlation between development time and fecundity. For particular cases the signs of the covariances between traits can be predicted. These predictions are tested using data drawn from the literature.     

Disease effects on reproduction can cause population cycles in seasonal environments

1. Recent studies of rodent populations have demonstrated that certain parasites can cause juveniles to delay maturation until the next reproductive season. Furthermore, a variety of parasites may share the same host, and evidence is beginning to accumulate showing nonindependent effects of different infections. 2. We investigated the consequences for host population dynamics of a disease-induced period of no reproduction, and a chronic reduction in fecundity following recovery from infection (such as may be induced by secondary infections) using a modified SIR (susceptible, infected, recovered) model. We also included a seasonally varying birth rate as recent studies have demonstrated that seasonally varying parameters can have important effects on long-term host-parasite dynamics. We investigated the model predictions using parameters derived from five different cyclic rodent populations. 3. Delayed and reduced fecundity following recovery from infection have no effect on the ability of the disease to regulate the host population in the model as they have no effect on the basic reproductive rate. However, these factors can influence the long-term dynamics including whether or not they exhibit multiyear cycles. 4. The model predicts disease-induced multiyear cycles for a wide range of realistic parameter values. Host populations that recover relatively slowly following a disease-induced population crash are more likely to show multiyear cycles. Diseases for which the period of infection is brief, but full recovery of reproductive function is relatively slow, could generate large amplitude multiyear cycles of several years in length. Chronically reduced fecundity following recovery can also induce multiyear cycles, in support of previous theoretical studies. 5. When parameterized for cowpox virus in the cyclic field vole populations (Microtus agrestis) of Kielder Forest (northern England), the model predicts that the disease must chronically reduce host fecundity by more than 70%, following recovery from infection, for it to induce multiyear cycles. When the model predicts quasi-periodic multiyear cycles it also predicts that seroprevalence and the effective date of onset of the reproductive season are delayed density-dependent, two phenomena that have been recorded in the field.     

Egg maturation dynamics of the parasitoid Microplitis rufiventris: starvation speeds maturation in early life

The number of mature eggs carried by a female parasitoid at any given moment (egg load) is a fitness‐related parameter affecting reproductive potential and impacting upon host population dynamics. Microplitis rufiventris Kokujev (Hymenoptera: Braconidae) is a solitary koinobiont endoparasitoid wasp of several noctuid pests, including Spodoptera littoralis. The number of mature eggs carried by females at emergence is approximately 50. The rate of egg maturation is strongly affected both by feeding status and access to host larvae. In early adult life, egg maturation rates are lower for 6–72 h in fed wasps compared with food‐deprived wasps. When given access to hosts, honey‐fed wasps live for approximately 9 days with high lifetime fecundity (226 eggs). By contrast to early adult life, the total realized fecundity is positively affected by feeding status, where water‐fed and starved females have 140 and 107 eggs, respectively. Egg resorption is most pronounced in the later life of females. The results suggest, in addition to confirming the effect of honey‐feeding on total fecundity, that fecundity of starved wasps includes rapid egg maturation early in life, which potentially could improve the performance of the parasitoid as a biological control agent.     

Seed predation,pathogen infection and life-history traits in Brassica rapa

Herbivory and disease can shape the evolution of plant populations, but their joint effects are rarely investigated. Families of plants of Brassica rapa (Brassicaceae) were grown from seeds collected in two naturalized populations in an experimental garden. We examined leaf infection by the fungus Alternaria, seed predation by a gall midge (Cecidomyiidae) and plant life-history traits. Plants from one population had heavier seeds, were more likely to flower, had less fungal infection, had more seed predation and were more fecund. Fungal infection score and seed predation rate increased with plant size, but large plants still had the greatest number of undamaged fruits. Spatial heterogeneity in the experimental garden was significant; seed predation rate and fecundity varied among blocks. An apparent tradeoff existed between susceptibility to disease and seed predation: plants with the highest fungal infection score had the lowest seed predation rate. Alternaria infection varied between populations, but the disease had no effect on fecundity. Seed predation did reduce fecundity. Damaged fruits had 31.4% fewer intact seeds. However, evidence for additive genetic variation in resistance to seed predation was weak. Therefore, neither disease nor seed predation was likely to be a strong agent of genetically based fecundity selection.     

The direct effects of male killer infection on fitness of ladybird hosts (Coleoptera: Coccinellidae)

Male killing bacteria are common in insects and are thought to persist in host populations primarily by indirect fitness benefits to infected females, whereas direct fitness effects are generally assumed to be neutral or deleterious. Here, we estimated the effect of male killer infection on direct fitness (number of eggs laid, as a measure of fecundity, together with survival) and other life‐history traits (development time and body size) in seven ladybird host/male killer combinations. Effects of male killers on fecundity ranged, as expected, from costly to neutral; however, we found evidence of reduced development time and increased survival and body size in infected strains. Greater body size in Spiroplasma‐infected Harmonia axyridis corresponded to greater ovariole number and therefore higher potential fecundity. To our knowledge, this is the first report of direct benefits of male killer infection after explicitly controlling for indirect fitness effects. Neutral or deleterious fitness effects of male killer infection should not therefore be automatically assumed.     

Experimental exposure of juvenile snails (Potamopyrgus antipodarum ) to infection by trematode larvae (Microphallus sp.): infectivity, fecundity compensation and growth

Host-parasite interactions that result in host castration are evolutionarily similar to predator-prey interactions because both interactions terminate reproduction for the host or prey. Yet, host-parasite interactions differ from predator-prey interactions in that infected hosts remain alive and potentially can make adjustments to their life-history strategy before castration is complete. Here we exposed juvenile snails (Potamopyrgus antipodarum) to infection by a digenetic trematode (Microphallus sp.) in order to determine whether: (1) pre-reproductive individuals could be infected, (2) individuals that were exposed to infection shifted resources to early reproduction (fecundity compensation), and (3) infected individuals exhibit altered growth rates relative to uninfected individuals. We found that juveniles are susceptible to infection; hence P. antipodarum could be selected for earlier maturation in populations where the risk of infection is high. We also found that fecundity compensation does not occur in this snail. Finally, we found that Microphallus-infected snails exhibit altered growth rates; individuals infected as juveniles have lower growth rates and are smaller than uninfected snails. These results suggest that growth is altered by infection of a trematode parasite but reproduction in uninfected snails is not induced by exposure to trematode eggs. Received: 11 January 1998 / Accepted: 19 May 1998     

Growth and survival effects on maturation pattern in populations of grayling with recent common ancestors

Mortality and growth rates were shown to influence maturation patterns in five populations of grayling ( Thymallus thymallus ) in central Norway. The populations share recent common ancestors as they derive from introductions performed in 1910, and they inhabit lakes with different environmental conditions (i.e. length of growth season, lake area and fishing pressure). Mortality rate (range of Z -values: 0.36–0.77) and growth pattern varied strongly among the populations. Mortality rates were negatively associated with population mean age at maturity ( r _sp=−0.90), supporting life-history theory which predicts early maturation to be favoured under conditions with high adult mortalities. Maturation reaction norms differed significantly among the populations. Individuals from one population showed no maturation plasticity (all individuals matured at age three), whereas rapid growers were found to mature earlier than slow growers in the remaining four populations. Life-history theory is again supported as it predicts rapid growers to mature early due to high age-specific fecundity and short generation times. Given low mortality risks, slow growers are predicted to delay maturation in order to gain high first-time fecundity. In high-mortality systems all individuals are predicted to mature early. This theory is supported by the present data as populationwise maturation plasticity increased with decreasing mortality rates. In the population with no maturation plasticity the corresponding high mortality rates were probably due to high fishing pressures.     

The function of multiple mating in oviposition and egg maturation in the seed beetle Callosobruchus maculatus

Abstract.  To comprehend the significance of multiple mating in female Callosobruchus maculatus , the material-benefits hypothesis is tested, as well as the potential function and mechanism of additional mating for females. The results show that longevity and lifetime fecundity are significantly higher in doubly-mated females, whereas females subject to an interrupted second copulation (which transfers no ejaculate) show resemblance with singly-mated females in lifetime fecundity and daily fecundity, supporting the material-benefits hypothesis. Female lifetime fecundity increases with ejaculate size in doubly-, but not singly-mated females. Doubly-mated females exhibit an immediate increase in fecundity on the day of remating. Moreover, dissection of ovaries after remating shows that the number of unlaid eggs is lower in doubly-mated females in comparison with singly-mated females, whereas the total number of unlaid and laid eggs shows no significant difference. This suggests that the second mating increases the oviposition rate, but not the egg maturation rate in female C. maculatus . Thus, ejaculate serves as a stimulus for oviposition, at least on the remating day. However, because doubly-mated females lay more eggs in total than singly-mated females, doubly-mated females appear to mature more eggs than the singly-mated counterparts. This is attributed to the 'indirect-driven' hypothesis, which states that the reduction in the number of mature eggs in the oviducts brought about by oviposition stimulates the maturation of oocytes in ovarioles to replenish the number of mature eggs.     

Changes in reproductive strategy in the ruffe during a period of establishment in a new habitat

The pattern of maturation, body size and fecundity was examined in a population of ruffe ( Gymnocephalus cernuus L.) three times during a period of rapid growth, and eventual stabilization, following its introduction to a new habitat. When the ruffe were less common, maturing ruffe were relatively large and immature ruffe relatively small, compared with when the ruffe were abundant. Intermediate ruffe population size showed a maturation pattern intermediate between these two extremes. It is suggested that this pattern of maturation is a response of the ruffe population to changing growth opportunity induced by changing intraspecific competition. This fluctuating maturation pattern is interpreted in terms of a threshold-dependent maturation trigger, operating on the rate of accumulation of energy and a trade-off between somatic growth and gonad development. When the ruffe population was large, high intraspecific competition resulted in low opportunity for growth; only fish with the highest rate of food acquisition were able to mature in a given year–the investment in gonadal tissue reducing somatic growth. When the ruffe population was low, the high rate of energy acquisition in the population resulted in the triggering of maturation, even at small size, only the very smallest fish remaining immature. High growth opportunity allowed maturing fish to develop gonad and maintain somatic growth. The pattern of size related fecundity also changed over the three periods. When growth opportunity was low, size related fecundity was greater than when opportunity for growth was high. This suggests that maturing females faced with poor growth conditions compensated by increasing egg number for a given body size either by decreasing egg size or by increasing total investment in ovarian tissue.     

Egg-eating age-structured predators in interaction with age-structured prey

In this paper, we consider a system of integrodifferential equations which models a predator-prey system with both species (predator and prey) age-structured and predators living only on the eggs of prey. The present model is a generalization of the model given in [20]. The existence, stability, and instability of nonnegative equilibria is studied assuming a general fecundity rate function for the prey. With a special choice of fecundity rate function for the predator it is shown here that a large maturation period m of the predator leads to stability. This seems to be contrary to the usual rule of thumb that increasing delays in growth rate responses cause instabilities.     

The regulation of mortality and fecundity in Schistosoma mattheei following a single experimental infection in sheep.

The regulation of mortality and fecundity of Schistosoma mattheei in sheep was examined using a series of mathematical models applied to data culled from the literature. Parasite mortality (μ) was found to be an increasing linear function of the magnitude of the initial infection over the ranges of doses examined (200–91, 000 cercariae) where μ = 9.78 × 10⁻³ + 3.476 × 10^−7* infection dose. Parasite fecundity (λ) was found to be inversely related to the duration of the infection. The best fit model for parasite fecundity was one in which fecundity decreased exponentially with time since initial infection, λ = λ₀e^−δ1−r. There was no evidence for density-dependent regulation of fecundity.     

Phosvitins in Fundulus oocytes and eggs. Preliminary chromatographic and electrophoretic analyses together with biological considerations

Vitellogenin serves as the plasma precursor for the yolk proteins, lipovitellin and phosvitin, in nonmammalian vertebrates. 32P-Vitellogenin was isolated from the plasma of the teleost, Fundulus heteroclitus, and was used both to label phosvitin in the ovary and to indicate the phosvitin region in preparative chromatographs of ovarian extracts on DEAE-cellulose. Crude [32P]phosvitin could be resolved further into two labeled components with shallow gradients on DEAE-cellulose and into eight labeled components by electrophoresis on 12% polyacrylamide gels. Only the two largest electrophoretically resolved components could be correlated with Coomassie Blue staining bands, but several of the smaller components could be indicated with the cationic carbocyanine dye, Stains-all. Stains-all-dyed components were also generally indicated as multiple bands. The ovary of a reproductively active female contains vitellogenic oocytes, postvitellogenic oocytes undergoing maturation prior to ovulation, and ovulated eggs. Examination of various types of follicles and eggs on polyacrylamide gels revealed that during maturation, the largest phosvitin components formed during vitellogenesis either disappear or diminish, while smaller phosvitin components appear. The transformation of phosvitin components can also be achieved in vitro by incubating prematurational follicles in a saline medium containing deoxycorticosterone. These preliminary results demonstrate that a complex array of phosvitin-like components are present within a single ovary of F. heteroclitus. We also postulate that one reason for the anomalous yolk proteins generally found thus far in teleost eggs is that some of the proteins derived from vitellogenin during vitellogenesis undergo further proteolysis during oocyte maturation.     

Elevated CO2 and tree fecundity: the role of tree size, interannual variability, and population heterogeneity

Long‐term population effects of changes in atmospheric CO₂ will be largely determined by reproductive effort. Our research objectives were to quantify variability in seed production and rate of maturation among individual Pinus taeda L. (Pinaceae) trees growing in elevated CO₂ (ambient plus 200 μL L^?1) since 1996. Estimating tree fecundity in nature is frustrated by the difficulty of counting seeds from individual trees and the need for long‐term data. We have used a hierarchical Bayes approach to model individual tree fecundity, accounting for the complexity of experimentation in a natural setting over multiple years. The study presented here demonstrates large variability in natural fecundity rates and contributes to our understanding of how both interannual variation and population heterogeneity influence elevated CO₂ effects. We found that trees growing under elevated CO₂ matured earlier and produced more seeds and cones per unit basal area than ambient grown trees. By 2004, trees grown in high CO₂ had produced an average 300 more seeds per tree than ambient grown trees. Although there was a trend toward decreasing mean CO₂ effect (difference in fecundity between elevated and ambient treatments) over time, the hierarchical analysis indicates that this decrease comes from the emergence of a few highly fecund ambient grown trees by 2002, rather than acclimation or downregulation among the fumigated trees. The most important effect of increased CO₂ in forest ecosystems may be the increase in fecundity reported here. Although biomass responses can sometimes be large, the increase in fecundity can have long‐term impacts on forest dynamics that transcend the current generation.     

Reproductive guilds (maturation,spawning frequency and fecundity) in the black pomfret,Parastromateus niger (Carangidae) in the Kuwaiti waters of the Arabian Gulf

Oogenesis, oocyte maturation pattern, spawning rhythm, spawning frequency, batch fecundity and oocyte diameter–frequency distribution of the black pomfret, Parastromateus niger (Bloch, 1795) in Kuwaiti waters were investigated from October 2003 to September 2005, using histological and morphological methods. The process of development is divided into four major phases: (i) primary growth phase; (ii) secondary growth phase; (iii) maturation phase; and (iv) spawning phase, followed by the regressed phase. Development of the yolky oocyte is an asynchronous process resulting, by the time of oocyte maturation, in a clear differentiation between a ready batch of oocytes (ready for spawning) and a reserve pool. Consequently, P. niger is capable of spawning multiple times throughout the reproductive season. Spawning frequency estimates, based on final oocyte maturation (FOM) method indicated that the species spawns once every 2.8 days during an 8‐month spawning season lasting from February to September, with a potential annual number of 22.4 spawns. Batch fecundity (BF) (2132–2001 648, mean 406 010 eggs), was significantly positively related to both standard length (SL) (P < 0.05) and ovary‐free body weight (OFBW) (P < 0.05), both parameters being good predictors of BF (r² = 30.8% for SL, from 22 cm onwards, and r² = 29.6% for OFBW, from 129.5 g onwards). No significant differences in monthly BF were found throughout the spawning season. Relative batch fecundity was 336 eggs/g OFBW; thus, estimate for potential annual relative batch fecundity was 7526 eggs g^?1 OFBW. The oocyte diameter–frequency distribution analysis revealed a multimodal distribution (at 100–200, 300–400 and 500–700 μm), confirming the evidence of multiple spawning.     

RELATIONSHIPS BETWEEN GENETIC VARIABILITY AND LIFE-HISTORY FEATURES OF BONY FISHES

Correlations between genetic variation and life-history variables were obtained for 80 species of bony fishes as a means of testing the hypothesis that genetic variation is directly related to 1) opportunity for balancing selection, as indicated by fecundity, and 2) environmental variation, as indicated by capacity for population increase. Genetic data were taken from the literature, and data on longevity, age at maturity, egg size, body size, and lifetime fecundity were taken from the literature where available and were otherwise estimated from other variables. Average heterozygosity does not increase significantly with increasing fecundity. However, heterozygosity is significantly associated with short generation times, quick maturation, small maximum size, and small eggs. Thus, heterozygosity appears to increase on a demographic continuum toward maximum values in species that are most strongly selected for maximizing the intrinsic rate of increase. Such species are associated with less stable environments. Thus, the results indicate a predominate role for environmental variation in controlling genetic variation of bony fishes.     

Resource-mediated demographic variation during the midsummer succession of a cladoceran community

SUMMARY. 1. The midsummer succession of cladoceran zooplankton in Lake Texoma was examined by demographic characterization of lake populations and a parallel set of life table experiments in which the dominant cladoceran species were raised on naturally occurring lake phytoplankton at lake temperatures. Diaphanosoma leuchtenbergianum occurred throughout the study period (mid June-mid August 1981) but Daphnia parvula, D. galeata mendotae and Ceriodaphnia lacustris were rarely detected after 15 July.
2. Analyses of size structure and size-specific fecundity of the cladoceran populations showed that (1) size structure shifted to smaller individuals in mid July, (2) size-specific fecundity was relatively constant in the smaller size classes and was constant or declined in larger females of each species, and (3) size at maturation did not vary significantly during the study of any population.
3. Individual growth rates of the two Daphnia species and Ceriodaphnia in the life table experiments were significantly lower in mid July when populations were declining and when size structure of each population was more dominated by smaller Individuals. The reduction in individual growth rates was short-lived in the life table experiments; growth rates returned to pre-decline levels by the end of July.
4. Two effects of temporary reductions in individual growth rates when size-specific fecundity and size at maturation are constant are (1) that size structure is temporarily shifted to smaller individuals, and (2) the potential rate of population growth is reduced because of delays in reproduction. Because data on size structure and size-specific fecundity are commonly used to evaluate the relative effects of predators and resource changes on populations it is noteworthy that erroneous conclusions about the mechanism of population decline may be reached when individual growth rates vary.     

Reproduction of Oligosarcus hepsetus (Cuvier, 1829) (Characiforms) in the Serra do Mar State Park, S?o Paulo, Brazil.

The purpose of this study was to characterize the reproduction of Oligosarcus hepsetus in the Grande river (Santa Virginia Unit - Serra do Mar State Park) based on the reproductive period, length of first gonadal maturation for both sexes, sexual proportion, oocyte development, type of spawn and fecundity. The Grande river was sampled monthly from January through December 2004. The numerical frequency of the stages of maturity and the volumetric method were utilized to determine the reproductive period and fecundity, respectively. The reproduction of Oligosarcus hepsetus was characterized by a long period, relatively low fecundity, precocious length of first gonadal maturation and parceled spawning.     

Cortisone-sensitive, innate resistance to Hymenolepis nana infection in congenitally athymic nude rats

The innate resistance of the unnatural rat host to the mouse tapeworm Hymenolepis nana is cortisone sensitive but thymus independent. When congenitally athymic nude rats were orally given eggs, cysticercoids, or adult worms of H. nana, no lumenal adults were established except when they were treated with cortisone acetate during the expected lumenal development. The effect of cortisone to promote adult maturation in the rats was compared in nude and normal rats given eggs of H. nana. The fecundity of the worms (assessed by the fresh worm biomass and the number of infective eggs produced) was much higher in cortisone-treated nude rats than in cortisone-treated normal rats. When the nude rats reconstituted with thymocytes were given eggs and treated with cortisone, the fecundity of H. nana dropped to the same level as in cortisone-treated normal rats. It is strongly suggested that the unnatural rat host has thymus-independent cortisone sensitive resistance to an initial infection (which is the main component of the innate resistance and blocks the lumenal establishment of this parasite) and thymus-dependent resistance (which suppresses the established worms' fecundity and may be ascribed to acquired resistance to the ongoing infection).     

Influence of host age and host deprivation on longevity, egg load dynamics and oviposition rate in Microplitis rufiventris

Adult size, longevity, egg load dynamics and oviposition ofMicroplitis rufiventris Kok. which began their development in the first, second, third （preferred hosts） or fourth （non-preferred hosts） instar larvae of Spodoptera littoralis （Boisd.） were studied. The parasitoid size was largely determined by the initial host size at parasitism. Non-ovipositing females derived from older hosts lived for longer periods than those derived from younger ones. However, the ovipositing females, irrespective of their size, lived for almost the same periods. At emergence, the oviducts of adult females contain a significant amount of mature eggs available for oviposition for a few hours on eclosion day. Egg load increases during the early phase of adult life. The amount of additional mature eggs and rate of egg maturation per hour was greater for wasps derived from preferred hosts compared with those in females derived from non-preferred hosts. The pattern of egg production in M. rufiventris females depended on the availability of hosts for parasitization. Host-deprived females depleted the egg complement with aging; the longer the host deprivation, the lower the oviduct egg load. Marked reduction in both realized or potential fecundity of host-deprived females was observed following host availability. Host privation for more than 3 days induced a marked deficit fecundity pattern through the female＇ s life. The realized fecundity was determined by the interaction among host availability, the number of eggs that are matured over the female＇ s life span, oviposition rate and host size from which the female was derived. These results suggest that： （i） M. rufiventris wasp is a weak synovigenic species; （ii） the maturation of additional eggs is inhibited once the maximum oviduct egg load is reached; （iii） the egg load of the newly emerged female is significantly less than the realized fecundity; and （iv） because M. rufiventris females oviposit fewer eggs when they begin depleting their egg supply at 3 days, augmentative releases will require release immediately following emergence to ensure the highest parasitization rate in the field.     

A field study using the fungicide benomyl to investigate the effect of mycorrhizal fungi on plant fitness

Summary The effect of vesicular-arbuscular mycorrhiza (VAM) on the fecundity ofVulpia ciliata ssp.ambigua was investigated at two field sites in eastern England by applying the fungicide benomyl to reduce VAM infection. The application of benomyl at the two sites produced very different results. At one site the application of the fungicide reduced the fecundity of plants whereas at the other fecundity was increased. At the first site the reduction in fecundity was linked to a significant reduction in VAM infection on the sprayed plants. The mechanism of the benefit associated with the VAM infection is however unclear: there was no treatment effect on morphology or on phosphorus inflow. At the second site, where fecundity was increased, there was only a negligible amount of VAM infection amongst the unsprayed plants and it is suggested that the increase in fecundity with the application of benomyl may have resulted from a reduction in infection by other, presumably pathogenic, fungi. The value of VAM fungi to the host plant may therefore not be restricted to physiological benefits. They may also provide protection to the plant by competing for space with other species of pathogenic fungi.