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

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

Life-history plasticity under resource stress in a clonal fish (Poeciliidae: Poeciliopsis)

Within-individual plasticity for reproductive investment was examined in a clonal fish (Poeciliidae: Poeciliopsis) grown under six levels of resource stress. Growth, age at first reproduction, egg production, egg size, egg energetic content, and survivorship were measured from fish grown in three dietary and two density treatments. Growth and fecundity decreased with both increased density and food stress. Age at first reproduction increased with increased density, but was unaffected by the diet treatments. Reproductive effort (clutch size per female weight), offspring investment (egg size and egg energetic content), and survivorship were invariant across all treatment combinations. We compare these results with predictions based on theoretical treatments of optimal reproductive investment.     

Monophyletic origin of multiple clonal lineages in an asexual fish (Poecilia formosa)

Despite the advantage of avoiding the costs of sexual reproduction, asexual vertebrates are very rare and often considered evolutionarily disadvantaged when compared to sexual species. Asexual species, however, may have advantages when colonizing (new) habitats or competing with sexual counterparts. They are also evolutionary older than expected, leaving the question whether asexual vertebrates are not only rare because of their 'inferior' mode of reproduction but also because of other reasons. A paradigmatic model system is the unisexual Amazon molly, Poecilia formosa, that arose by hybridization of the Atlantic molly, Poecilia mexicana, as the maternal ancestor, and the sailfin molly, Poecilia latipinna, as the paternal ancestor. Our extensive crossing experiments failed to resynthesize asexually reproducing (gynogenetic) hybrids confirming results of previous studies. However, by producing diploid eggs, female F(1) -hybrids showed apparent preadaptation to gynogenesis. In a range-wide analysis of mitochondrial sequences, we examined the origin of P. formosa. Our analyses point to very few or even a single origin(s) of its lineage, which is estimated to be approximately 120,000 years old. A monophyletic origin was supported from nuclear microsatellite data. Furthermore, a considerable degree of genetic variation, apparent by high levels of clonal microsatellite diversity, was found. Our molecular phylogenetic evidence and the failure to resynthesize the gynogenetic P. formosa together with the old age of the species indicate that some unisexual vertebrates might be rare not because they suffer the long-term consequences of clonal reproduction but because they are only very rarely formed as a result of complex genetic preconditions necessary to produce viable and fertile clonal genomes and phenotypes ('rare formation hypothesis').     

The adaptive value of functional and life-history traits across fertility treatments in an annual plant

Background and Aims

Plant functional traits are assumed to be adaptive. As selection acts on individuals and not on traits, interpreting the adaptive value of a trait not may be straightforward. For example, productive leaves are associated with fertile environments. However, it is not clear if productive leaves confer an advantage in these habitats, or if they are an advantage as part of a suite of coordinated traits.

Methods

Genotypes of Arabidopsis thaliana were grown in high and low nutrient treatments and low, neutral and high pH treatments. Nutrient availability is reduced in acidic or basic soils relative to neutral pH soils. pH treatments were used to alter the availability of resources rather than the amount of resources.

Key Results

Leaf function (specific leaf area, SLA) and life history (size at reproduction, age at reproduction) were variable across genotypes and were plastic. High nutrient availability induced higher SLA and larger size at reproduction. Genotypes that reproduced at large size in high nutrient conditions at neutral pH had the greatest fruit production. SLA was only indirectly related to fruit production through a causal relationship with rosette size; in high nutrient conditions, plants with high SLA were large at reproduction and had higher fruit production. In high nutrient and high pH treatments, plants were large at reproduction, but large size at reproduction was associated with low fecundity. This suggests that large size is adaptive under high nutrient availability.

Conclusions

Interpreting the adaptive value of functional traits will sometimes only be possible when these traits are considered as a suite of correlated and coordinated traits. Leaf functional traits may be important in defining adaptive strategies in A. thaliana but only through how they affect plant life history. Finally, manipulating soil pH can be a valuable tool in assessing adaptive plasticity on nutrient gradients.     

Trade-offs between sexual and clonal reproduction in an aquatic plant: experimental manipulations vs. phenotypic correlations

That trade-offs result from the allocation of limited resources is a central concept of life history evolution. We quantified trade-offs between sexual and clonal reproduction in the aquatic plant, Butomus umbellatus, by experimentally manipulating sexual investment in two distinct nutrient environments. Increasing seed production caused a significant but nonlinear trade-off. Pollinating half of all flowers strongly reduced clonal bulbil production, but pollinating the remaining flowers did not cause any further trade-off. Trade-offs were not stronger under low nutrient conditions that clearly limited plant growth. Experimentally induced trade-offs were not reflected in negative phenotypic correlations between sexual and clonal allocation among plants within eight populations grown in a uniform greenhouse environment. Diminishing effects of increased sexual allocation plus a lack of accord between experimental manipulations and phenotypic correlations suggest that trade-offs between sexual and clonal reproduction are unlikely to constrain the evolution of reproductive strategy in this species.     

Developmental instability in Brassica campestris (Cruciferae): fluctuating asymmetry of foliar and floral traits

The degree of fluctuating asymmetry of bilateral traits provides a measure of developmental instability, which can be influenced by genetic as well as environmental stress. We studied genetic variation between and within two populations of the mustard Brassica campestris for asymmetry of foliar (cotyledon width) and floral (petal length and width) traits as well as for phenological (germination and flowering) and performance (biomass and flowering) traits. The two populations differed in mean expression of most traits, including asymmetry. However, within-population estimates of genetic variability tended to be lower for asymmetry than other traits. Asymmetry was greater in the population that had lower biomass accumulation and flower production, which supports the idea that population-level asymmetry may be indicative of population-level performance. However, within each population, evidence that performance was negatively correlated with asymmetry was equivocal. Within populations there was little or no concordance among estimates of asymmetry based on different structures, i.e., plants that had highly asymmetrical cotyledons did not tend to have highly asymmetrical petals. The lack of a general buffering capacity at the individual level may be explained by developmental processes (e.g., action of different genes or morphogens) as well as evolutionary processes (e.g., selection on asymmetry of different traits).     

Phenotypic plasticity in life-history traits of femaleThalassoma bifasciatum (Pisces: Labridae): 2. Correlation of fecundity and growth rate in comparative studies

Synopsis The cost of reproduction is a central concept in theories of life-history evolution. One way to empirically examine the tradeoff between current reproduction and future reproductive prospects is to use natural intraspecific variation in life-history traits. However, this approach is complicated by the sensitivity of life-history traits to variation in the level of resources. We report here an attempt to measure the cost of increasing reproductive activity in populations of female bluehead wrasse,Thalassoma bifasciatum, a coral-reef fish. All of the significant correlations of fecundity and growth rate were positive, in contradiction to the tradeoff predicted by the cost concept. In one of two regions studied, the populations with relatively high mean growth rate had a relatively large mean fecundity. The trait means were also positively associated over time: in months of rapid growth, female reproductive activity was high. Even after removing the effects of habitat and time period in a comparison of individual traits, no growth cost to reproduction appears. Variation in the abundance of resources over space and time is likely to interfere with the measurement of the cost of reproduction in many natural systems.     

Seasonal changes in the reproductive and life-history traits of Chrysomela populi L. (Coleoptera: Chrysomelidae)

Seasonal changes in the reproductive and life‐history traits of Chrysomela populi L. were investigated in Shobara (34°52′N, 133°01′E), Japan, in 2004 and 2005. Larvae were hatched and reared under natural photoperiod and temperature in 8 different periods between late April and early September in 2004. The incidence of adult diapause increased with progression of the season in 2004. The critical timing of diapause induction, as defined by 50% of females entering diapause, was estimated to occur between late July and early August. No effects of season on the survival rate from hatching to adult emergence, percentage ovipositing females and percentage females that deposited viable eggs were clearly detected. Adult weight at emergence fluctuated seasonally, which was probably caused by air temperature. No effect of season on the oviposition period was observed among females that averted diapause. Females that entered diapause in 2004 started oviposition from the first day of May in 2005. Reproductive output was significantly greater in diapause females than in non‐diapause females. This increase in the reproductive output of diapause females was caused by elongation of the oviposition period and consequently by enlargement of the number of egg clutches deposited. These results suggest that the seasonal timing of diapause induction in females of C. populi would be affected by various risks and benefits to reproduction and survival.     

Size at the onset of sexual maturity in the anomuran crab, Aegla uruguayana (Aeglidae)

The size at maturity was studied in the crab Aegla uruguayana from the Areco River (31°14′ S, 59°28′ W), Argentina. Size at sexual maturity was determined according to three criteria: morphometric (change in the relative growth of reproductive characters), histological (first maturation of gonads) and functional (capability to mate and carry eggs). Regarding females, morphometric maturity occurred at a carapace length (CL) of 11.50 mm, considering abdomen width as a reproductive character. Gonad maturity of females could be observed at a minimum size ranging from 15 to 17 mm CL. The smallest ovigerous female observed in the field was 15.60 mm CL, although a relevant population incidence of ovigerous females (86.6%) has just been observed at values higher than 17 mm CL. As for males, the relative growth of the left chela length changed at a value of 15.40 mm CL, while morphological changes in sexual tube occurred between CL of 14 and 16 mm. Testicular maturation occurred at a CL ranging from 17 to 19 mm. The smallest size of males having spermatozoids in their vasa deferentia was 18.70 mm CL. The results obtained indicated that, in both sexes, functional maturity occurred after morphometric maturity and at a size similar to that of gonad maturity. Comparing sexes, females acquired sexual maturity (morphometric, gonad and functional maturity) at sizes statistically smaller than those of males.     

Vegetative reproduction and clonal diversity in Rhytidium rugosum (Rhytidiaceae, Bryopsida) inferred by morpho-anatomical and molecular analyses

Many bryophytes exhibit specific life forms that result in the formation of distinct patches. This is primarily achieved by consequent vegetative multiplication through indeterminate growth and vegetative reproduction. Little, however, is known about the underlying mechanisms and genetic patterns. In this study on vegetative multiplication in bryophytes, we apply morpho-anatomical and molecular (AFLP fingerprinting) techniques to analyze the vegetative reproduction system, clonal diversity, and habitat colonization of a pleurocarpous moss, Rhytidium rugosum (Rhytidiaceae). Morpho-anatomically, three types of vegetative diaspores are identified and illustrated: ramets, separated after decay and disintegration of older shoot parts (clonal reproduction); brood branches/branchlets; and caducous shoot apices (both vegetative reproduction s.str.). The AFLP fingerprinting of 35 samples (from two plots in Thuringia, further German populations, and from France, Russia, and Canada) resulted in the identification of three clones from the two plots, each comprising two to 15 samples with identical fingerprints. Samples from one clone occurred in both plots, thus proving dispersal of vegetative diaspores. The AFLP analysis further revealed close relationships of the plot samples, which suggest a clonal rather than generative (sexual) origin of the population.     

Relationships of reproductive traits and body size with attainment of sexual maturity and age in Blanding's turtles (Emydoidea blandingi)

To place associations among body size, age at maturity, age, and reproductive traits of a long-lived organism in the context of current life history models based on the concept of norms of reaction, we examined data from a mark-recapture study of Blanding's turtles (Emydoidea blandingi) in southeastern Michigan during 24 of the years between 1953 and 1988. Females matured between 14 and 20 years of age. Both the smallest and largest adult females in the population were reproducing for the first time in their lives. This result suggests that a combination of differences in juvenile growth rates and ages at maturity, and not indeterminate growth, are the primary cause of variation in body size among adults. Body size variation among individuals was not related to age at sexual maturity. Females that had slower growth rates as juveniles matured later at similar mean body size compared to those with more rapid growth that matured at an earlier age. As a result, a linear model of age at sexual maturity with growth rates of primiparous females between hatching and maturity was significant and negative (R² = 0.76). Frequency of reproduction of the largest and smallest females was not significantly different. Clutch size did not vary significantly with age among either primiparous or multiparous females. Clutch sizes of primiparous females and multiparous females were not significantly different. However, older females (>55 years minimum age) reproduced more frequently than did younger females (minimum age <36 y).     

Population biology and size at the onset of sexual maturity of the amphidromous prawn Macrobrachium olfersii (Decapoda,Palaemonidae) in an urban river in southeastern Brazil

Abstract

Information on population biology is essential for a better understanding of the ecological stability of a species and the ecosystem in which it lives, especially in areas affected by anthropogenic activities. This study investigated the population biology of Macrobrachium olfersii in an ecosystem under human impact, focusing on sex ratio, reproductive period, and juvenile recruitment. The onset of sexual maturity was also estimated. Samplings were performed monthly in the daytime from June 2006 to May 2007 on the Taquaral River in the city of Ubatuba, São Paulo State, Brazil. The sex ratio was skewed towards females, and both the reproductive period and the juvenile recruitment were continuous. The estimated size at which the species reached sexual maturity corroborated studies on populations from other locations. Results from the current study strongly indicate that the ecosystem studied may have suffered environmental stress, which probably interfered with the migration and establishment of the species: during the study, the species was found in significant abundance, but six years later, it was not captured in samplings.     

Differences in life-history and aging in two clonal groups of Daphnia cucullata Sars (Crustacea: Cladocera)

Life-history traits were compared in twenty-two clones of Daphnia cucullata from the pre-alpine lake Klostersee in southern Germany. Clones were kept under standardized conditions (18 °C; 16:8 L/D) both singly and with 25 females per glass. Clones belonged to two genotype groups distinguished by enzyme electrophoresis. 11 of the clones had the genotype FM at the phosphate glucose isomerase locus and 11 the genotype MM. Neonates and adults of the age class 10 to 21 days were larger in one group (FM) than in the other group (MM). The size specific growth rates within any age class depended on age and body size and differed between the genotype groups. The fecundity of young FM animals (< 16 days) was significantly higher than that of MM animals, but for older females fecundity did not differ significantly. Senescence started about 6 days earlier in FM than in MM. The data are consistent with results of a field study on Daphnia cucullata in the Klostersee (Jacobs, 1990).     

Growth and reproductive characteristics in artificially formed clonal gametophytes ofDryopteris filix-mas (Dryopteridaceae)

Artificially formed clonal gametophytes ofDryopteris filix-mas (L.)Schott were able to grow and reproduce, although growth rates, proportions of hermaphrodites and reproductive efforts were low. Variable density and nutrient levels appeared to affect gametophytic life histories and they continued to influence the viability of the developing sporophyte generation. Differences between populations and sporophytes were discovered in life histories. Hermaphroditic gametophytes were considerably larger and possessed higher viability than did males. Clear differences between clones in the level of phenotypic plasticity in gametophyte size were observed. No neighbor effect on gametophyte sex expression was detected with the densities and nutrient levels used.     

Effects of nutrient availability on primary sexual traits and their response to selection in Spergularia marina (Caryophyllaceae)

Abstract Theoretical models of the evolution of sex allocation generally assume a negative genetic correlation between components of male and female investment or function. To test this assumption, and to determine whether the expression of the correlation is sensitive to environmental conditions, we conducted an artificial selection experiment targeting primary sexual traits in the autogamous herb, Spergularia marina. Mass selection favouring individuals with high (or low) pollen and ovule production per flower had previously been conducted for two generations under uniform conditions. Following a third episode of selection (reported here) within the high‐pollen‐, high‐ovule‐, low‐pollen‐ and low‐ovule‐producing lineages, selected maternal seed families were replicated and cultivated in three nutrient treatments. With this design, we observed the effects of nutrient availability on: floral phenotype, the realized response to selection and the expression of genetic variation in, and covariation between, male and female investment. The first two episodes of selection Mazer et al. 1999 (Evolution 53:717–731) detected evidence for a genetically based negative correlation between ovule and anther production. Following the third episode of selection, phenotypic differences among the selected lines were maintained in all treatments, but evidence for a negative correlation between male and female investment nearly disappeared. Only under low‐nutrient conditions, in which plants selected for low ovule production exhibited elevated anther production, was a genetic trade‐off expressed. Either genetic drift or the combination of novel growing conditions (relative to previous generations) and G × E interactions may have caused the change in the expression of the genetic correlation. This experiment also allowed us to test the prediction that autogamous selfers should exhibit higher canalization of the ratio of male to female investment than of its components. Supporting this prediction, the A : O ratio was more strongly buffered against environmental variation than either anther or ovule production per flower.     

Reproductive performance of clonal and sexual bark beetles (Coleoptera: Scolytidae) in the field

In Ips acuminatus (Gyll.) parthenogenetic females occur together with sexual females and with sexual males upon which they depend for sperm. In a reciprocal‐transplant experiment, I studied fecundity differences among parthenogenetic and sexual females from two populations that differ dramatically in the proportion of clonal females. In a second experiment, I studied competition between larvae from different mothers and between females from the two source populations. Fecundity measured by the number of eggs per egg tunnel was influenced by the ambient environment at the sites of the experiment as well as the origin of the female, and was generally higher for clonal than for sexual females at both sites. In experimental groups where larvae competed with larvae from their own population (pure treatments), the number of surviving pupae was significantly lower than in groups where females from the two source populations were mixed. The high fecundity of clonal females makes coexistence of the two types of females difficult to explain. It makes the reproductive advantage associated with clonality in I. acuminatus even higher than the two‐fold difference due to asexuality per sé. The significant differences in the number of pupae in mixed vs. pure groups suggest ecological divergence between sexual and clonal females. This would make the mortality of larvae not only density dependent, but also frequency dependent, which could explain the coexistence of sexual and clonal females.     

Maternal allocation strategies and differential effects of yolk carotenoids on the phenotype and viability of yellow-legged gull (Larus michahellis) chicks in relation to sex and laying order

Egg quality may mediate maternal allocation strategies according to progeny sex. In vertebrates, carotenoids have important physiological roles during embryonic and post-natal life, but the consequences of variation in yolk carotenoids for offspring phenotype in oviparous species are largely unknown. In yellow-legged gulls, yolk carotenoids did not vary with embryo sex in combination with egg laying date, order and mass. Yolk lutein supplementation enhanced the growth of sons from first eggs but depressed that of sons from last eggs, enhanced survival of daughters late in the season, and promoted immunity of male chicks and chicks from small eggs. Lack of variation in egg carotenoids in relation to sex and egg features, and the contrasting effects of lutein on sons and daughters, do not support the hypothesis of optimal sex-related egg carotenoid allocation. Carotenoids transferred to the eggs may rather result from a trade-off between opposing effects on sons or daughters.     

Costs of reproduction in Nyssa sylvatica: sexual dimorphism in reproductive frequency and nutrient flux

Summary We examined the influence of differential reproductive frequency between the sexes on tertiary (phenotypic) sex ratios in the the dioecious tree Nyssa sylvatica (Nyssaceae). Reproduction was evaluated in relation to sex, size and canopy exposure using flowering data collected from 1229 marked trees over a four year period. For subsets of each population we used data on flower number, fruit crop size, fruit/flower ratios, and individual flower and fruit mass to compare biomass invested in reproductive structures of males and females. We also examined seasonal changes in stem nitrogen and soluble carbohydrate content in relation to flower and fruit production for trees of each sex. Our results indicate that: 1) Male-biased tertiary sex ratios could be explained by more frequent reproduction by male trees; 2) Estimated secondary sex ratios based on sums of all known males and females were not significantly different from 1:1; 3) Flowering frequency of males and females was significantly related to plant size (DBH) and exposure of the canopy to light; 4) Estimtes of reproductive biomass allocation ranged from 1.36 to 10.8 times greater for females relative to males; 5) Flower production was related to stem nutrient status for both sexes, but nutrient depletion and its effect on subsequent flowering was much more pronounced for female trees. We conclude that less frequent flowering by female trees may result from depletion of stored reserves, and that differential flowering frequency in N. sylvatica may ultimately reduce apparent sexual differences in the costs of reproduction.