FITNESS CONSEQUENCES OF VARIATION IN EGG SIZE AND FOOD ABUNDANCE IN BROOK TROUT SALVELINUS FONTINALIS

Maternal effects in the American coot: consequences for offspring growth and survival

Evolution of Egg Size in Free-Spawners: Consequences of the Fertilization-Fecundity Trade-Off

To test relationships between maternal size, egg size and size of new offspring, we studied (a) the effect of maternal size on egg size and number, and maternal survival after oviposition and (b) the effect of egg size on the duration of development and new imago size in three flea species (Xenopsylla ramesis, Synosternus cleopatrae, Parapulex chephrenis) with varying host and habitat specificity. In general, the number and size of eggs as well as total egg volume appeared to be independent of maternal body size. There was no trade-off between egg number and size. However, female body size was related to post-oviposition survival, with larger females surviving longer after oviposition than smaller females. In addition, females that produced more eggs died faster after oviposition in X. ramesis but not in the two other species. There were no significant size differences between eggs that developed into new imagoes and eggs that did not survive. Survivorship of male and female eggs did not differ; however, new adult females were significantly larger than new adult males. Female, but not male, new imagoes exhibited a significant positive relationship between egg size and duration of preimaginal development in all three species, with larger eggs developing faster than smaller eggs. In X. ramesis and S. cleopatrae, faster developing eggs also developed into larger new imagoes. We conclude that these patterns were largely consistent among the three flea species, suggesting that they result from the same mechanisms and are weakly affected by the ecological specialization of a given species.

The optimal balance of reproductive effort between offspring size and number depends on the fitness of offspring size in a particular environment. The variable environments offspring experience, both among and within life-history stages, are likely to alter the offspring size/fitness relationship and favor different offspring sizes. Hence, the many environments experienced throughout complex life-histories present mothers with a significant challenge to optimally allocate their reproductive effort. In a marine annelid, we tested the relationship between egg size and performance across multiple life-history stages, including: fertilization, larval development, and post-metamorphosis survival and size in the field. We found evidence of conflicting effects of egg size on performance: larger eggs had higher fertilization under sperm-limited conditions, were slightly faster to develop pre-feeding, and were larger post-metamorphosis; however, smaller eggs had higher fertilization when sperm was abundant, and faster planktonic development; and egg size did not affect post-metamorphic survival. The results indicate that egg size effects are conflicting in H. diramphus depending on the environments within and among life-history stages. We suggest that offspring size in this species may be a compromise between the overall costs and benefits of egg sizes in each stage and that performance in any one stage is not maximized.

1. Eggs from a layer-type breeder flock (Baladi, King Saud University) between 50 and 63 weeks of age were used in three trials to study the effects of electrical field (EF) during incubation on albumen and yolk heights, incubation temperature, egg weight loss and hatchability traits. The effects of egg size and eggshell characteristics on hatchability traits of eggs incubated under EF were investigated. 2. Eggs were weighed and graded into three weight classes (small, medium, and large). The physical dimensions, eggshell characteristics, and conductance of eggs were examined. The incubator was divided into two compartments for the control and EF treatments. Two aluminium plates were fitted on the inside walls of the EF compartment, face to face, and connected to a step up electric transformer. Eggs were exposed constantly to the EF during the first 18 d of incubation at the level of 30 kV/m, 60 Hz. 3. Egg size influenced the physical dimensions and eggshell characteristics of eggs. Large eggs had higher egg weight, egg surface area, egg volume, eggshell conductance, and eggshell weight and lower yolk weight percentage than medium or small size eggs. Small eggs had lower egg length and higher egg density than large or medium size eggs. Large eggs had higher eggshell thickness than small size eggs. 4. EF incubation of eggs raised incubation temperature by 0·06°C, and increased the percentage of egg weight loss, hatchability, and weight of hatching chicks and reduced the early embryo deaths, and length of incubation by approximately 9·8, 19·6, 1·7, 62·1 and 2·1%, respectively. 5. There was no significant difference between the two incubation treatments in the heights of albumen and yolk of incubated eggs, percentages of late embryo deaths, and pips with live and dead embryos. Hatchability traits were not significantly influenced by egg size. 6. It was concluded that EF incubation of eggs increased hatchability, chick-hatching weight, and reduced the length of incubation of Baladi eggs. Differences in the physical dimensions and eggshell characteristics of eggs did not influence hatchability traits of eggs under EF incubation.

Effects of egg size reductions on development time and juvenile size in three species of echinoid echinoderms: Implications for life history theory

This laboratory study of Daphnia magna had two major aims: (1) to examine the effect of food quantity on offspring size and number and how this effect varies with genotype and maternal size, age, and metabolic demand; and (2) to examine the relationship of investment per offspring to total reproductive investment. Differences in the response of offspring size to food quantity between the two clones in this study and between Daphnia studies in the literature are explained by nonlinear model that explicitly considers the effect of food quantity relative to the metabolic demands of the mother. According to this model, offspring size covaries positively with food quantity/maternal demand at very low food levels because of reproductive constraints, but it covaries negatively with food quantity/maternal demand at middle food levels as an adaptive response, and it remains constant at very high food levels because of a lower limit on viable offspring size. Effects of genotype (clone) and ration level on trade—offs between egg size and number and on the maternal size—dependence of egg mass, brood mass, and brood size are described. Strong genotype—environment interactions are evident for egg size and number. Trade—offs between egg size and number are seen at high low ration levels, but only in second (or third) to sixth broods, and not in first broods, which had relatively high variation in total mass. Results of this study that contradict predictions or assumptions of optimal offspring investment theory include: (1) at both ration levels both egg size and number were strongly positively correlated with brood mass, independently of maternal mass, (2) although, at low ration, variability of brood mass usually exceeded the variability of egg mass, as expected, the opposite was true at high ration, and (3) the interclonal difference in offspring size was not correlated with survival ability under starvation. Offspring size covaried positively with maternal size and age, partly independently of culture conditions. This relationship is explained as the result of selection in growing parthenogenetic populations for increased fecundity in young small females, which are constrained to produce relatively small eggs because of spatial limitations of the brood pouch.

A large number of studies have reported a positive relationship between the egg size of birds and the subsequent growth and/or survival of nestlings, but such effects may partly be due to confounding variables, e.g. parental quality. In order to evaluate the potential effects of egg size, and of parental quality, on early nestling growth in the Antarctic petrel, we performed an experiment in which eggs of different size were swapped between nests. From a sample of 300 nests with eggs of known size, we selected eggs belonging to the lower quartile (small eggs), and those belonging to the upper quartile (large eggs), with respect to volume. Half of the small eggs were exchanged with small eggs from other nests, and the other half with large eggs. A similar procedure was used for large eggs. Growth and survival of the nestlings were recorded until 12 days old. Hatching success was positively related to egg size. Egg size influenced nestling body mass until the age of 3 days, and tarsus length was affected until 12 days old. However, these effects were not due to an effect of egg size on growth rates, but reflected instead the influence of egg size on hatchling size. In contrast to most previous studies, we found no effect of parental quality (as reflected in the size of own eggs) on foster nestling size or growth until 12 days old. This could be because egg size does not reliably reflect parental quality in the species, or because parental effects become evident only at later nestling stages. We discuss why egg size variation is maintained in this and other species where egg size influences parental fitness through the survival of eggs or nestlings.

1. Maternal effects have been suggested to function as a mechanism for transgenerational plasticity, in which the environment experienced by the mother is translated into the phenotype of the offspring. In birds and other oviparous vertebrates where early development is within the egg, mothers may be able to improve the viability prospects of their offspring at hatching by priming eggs with immunological and nutritional components. 2. We studied how resource availability affects maternal investment in offspring by feeding Ural owl (Strix uralensis, Pall.) females prior to egg-laying in 3 years of dramatically different natural food conditions. 3. Supplementary feeding prior to laying increased body mass and the level of Igs of females measured at clutch completion. Supplementary fed Ural owl females laid larger eggs than control females, and had offspring with higher levels of Igs at hatching compared to offspring of control females. 4. We found variation in maternal allocation of resources to the eggs in response to environmental conditions: during a year of rapidly declining food abundance, maternal Igs in hatchlings were higher, whereas egg size was smaller compared to years with a more stable food supply. 5. Egg size had a positive effect on offspring body mass at fledging, whereas Igs at hatching did not affect Igs at fledging. 6. We conclude that maternal body condition and maternal Igs, as well as hatchling Igs and egg size are limited by food resources during egg production. Hatchlings rely on maternally derived Igs and, hence, our results suggest that mothers with high levels of Igs passively transfer higher Igs levels to their eggs instead of active manipulation of Igs levels in eggs. Ural owl egg size appears to be highly sensitive to short-term changes in food abundance, with important consequences for nestling growth.

Understanding the relationship between egg size, development time, and juvenile size is critical to explaining patterns of life-history evolution in marine invertebrates. Currently there is conflicting information about the effects of changes in egg size on the life histories of echinoid echinoderms. We sought to resolve this conflict by manipulating egg size and food level during the development of two planktotrophic echinoid echinoderms: the green sea urchin, Strongylocentrotus droebachiensis and the sand dollar, Echinarachnius parma. Based on comparative datasets, we predicted that decreasing food availability and egg size would increase development time and reduce juvenile size. To test our prediction, blastomere separations were performed in both species at the two-cell stage to reduce egg volume by 50%, producing whole- and half-size larvae that were reared to metamorphosis under high or low food levels. Upon settlement, age at metamorphosis, juvenile size, spine number, and spine length were measured. As predicted, reducing egg size and food availability significantly increased age at metamorphosis and reduced juvenile quality. Along with previous egg size manipulations in other echinoids, this study suggests that the relationship between egg size, development time, and juvenile size is strongly dependent upon the initial size of the egg.

Effects of Egg Size and Density on Metamorphic Traits in Tadpoles of the Natterjack Toad (Bufo calamita)

We investigated the relationship between egg size and composition and their subsequent effects on hatching and fledging success in Eurasian Oystercatchers Haematopus ostralegus on the island of Schiermonnikoog (53d̀30'N, 06d̀10'E) in the Dutch Wadden Sea between 1986 and 1990. Egg size variation was considerable. The ratio in egg volume of the largest over the smallest egg was more than 1.5 in each of five years. Differences between females accounted for, on average, 61% of the total variance of egg volume. Individual females produced similarly sized eggs from one year to another. Nevertheless, average egg volume per clutch declined by 0.2 cm3 per year as females aged, but this explains little of the observed variation. Female size and food supplementation had no significant effect on egg volume. In absolute terms, large eggs contain more lean dry matter and lipid than small eggs, but the proportion of both constituents decreases with egg size. Consequently, the combustible energy content of eggs increases less than proportionally with egg size. Hatchability averaged 87% and was not correlated with egg volume. Hatchling weight increased with egg size, averaging 65% of fresh egg weight. Chick survival until fledging did not increase with egg volume. Consequently, within the size range observed, large and small eggs are of comparable quality. Since both the costs and benefits of large eggs compared with small eggs seem small at best, we propose that, within the size range encountered in this population, egg size can be considered an evolutionarily neutral trait.

Optimal offspring size theory states that natural selection should balance reproductive output by optimizing between offspring size and offspring number. If a species has evolved an optimal offspring size, the fitness of larger females should be increased by simply producing more offspring of an optimum size. In contrast, when offspring size is not optimized, the morphological constraint hypothesis may apply, and in this case, maternal fitness is increased by producing the greatest number of the largest offspring that mothers are physically capable of producing. We used a log-log allometric regression approach on clutch size, egg size, and body size data to test the application of optimal offspring size theory and the morphological constraint hypothesis in the Mexican mud turtle (Kinosternon integrum) in southern Mexico. Our results indicate that this turtle seems to follow the morphological constraint hypothesis when all data are analyzed together, but when data are divided between small (< 140 mm plastron length) and large females (> 140 mm plastron length), optimal offspring (egg) size theory was supported only in large females, while the morphological constraint hypothesis was supported in small females. Our results thus indicate that K. integrum females may increase their fitness in two different, size-dependent ways as they grow from size at sexual maturity to maximum body size.

I manipulated egg size and followed individual mass trajectories from the egg stage in Atlantic salmon to test for effects of size, and for interactions between size and paternal body mass, on offspring performance in strongly food-limited environments. Egg size had a strong effect on body mass at yolk absorption, causing juveniles originating from large eggs to outgrow their siblings from small eggs. This corroborates previous findings of egg size effects under more benign environments, and demonstrates that positive effects of egg size on offspring success are manifested even under strong food-limitation. Previously reported negative effects of being large during the critical period for survival in dense populations are thus likely related to social interactions, rather than to effects of density on total food abundance in the environment. The effect of egg size on offspring performance, and hence the optimal egg size, was independent of paternal body mass.

1. Maternal reproductive investment is thought to reflect a trade-off between offspring size and fecundity, and models generally predict that mothers inhabiting adverse environments will produce fewer, larger offspring. More recently, the importance of environmental unpredictability in influencing maternal investment has been considered, with some models predicting that mothers should adopt a diversified bet-hedging strategy whilst others a conservative bet-hedging strategy. 2. We explore spatial egg size and fecundity patterns in the freshwater fish southern pygmy perch (Nannoperca australis) that inhabits a diversity of streams along gradients of environmental quality, variability and predictability. 3. Contrary to some predictions, N.australis populations inhabiting increasingly harsh streams produced more numerous and smaller eggs. Furthermore, within-female egg size variability increased as environments became more unpredictable. 4. We argue that in harsh environments or those prone to physical disturbance, sources of mortality are size independent with offspring size having only a minor influence on offspring fitness. Instead, maternal fitness is maximized by producing many small eggs, increasing the likelihood that some offspring will disperse to permanent water. We also provide empirical support for diversified bet-hedging as an adaptive strategy when future environmental quality is uncertain and suggest egg size may be a more appropriate fitness measure in stable environments characterized by size-dependent fitness. These results likely reflect spatial patterns of adaptive plasticity and bet-hedging in response to both predictable and unpredictable environmental variance and highlight the importance of considering both trait averages and variance. 5. Reproductive life-history traits can vary predictably along environmental gradients. Human activity, such as the hydrological modification of natural flow regimes, alters the form and magnitude of these gradients, and this can have both ecological and evolutionary implications for biota adapted to now non-existent natural environmental heterogeneity.