Adaptive plasticity in egg size helps butterflies to cope with desiccation.

Egg size is a widely-studied trait and yet the causes and consequences of variation in this trait remain poorly understood. Egg size varies greatly within many avian species, with the largest egg in a population generally being at least 50% bigger, and sometimes twice as large, as the smallest. Generally, approximately 70% of the variation in egg mass is due to variation between rather than within clutches, although there are some cases of extreme intra-clutch egg-size variation. Despite the large amount of variation in egg size between females, this trait is highly consistent within individuals between breeding attempts; the repeatability of egg size is generally above 0.6 and tends to be higher than that of clutch size or laying date. Heritability estimates also tend to be much higher for egg size (> 0.5) than for clutch size or laying date (< 0.5). As expected, given the high repeatability and heritability of egg size, supplemental food had no statistically significant effect on this trait in 18 out of 28 (64%) studies. Where dietary supplements do increase egg size, the effect is never more than 13% of the control values and is generally much less. Similarly, ambient temperature during egg formation generally explains less than 15% of the variation in egg size. In short, egg size appears to be a characteristic of individual females, and yet the traits of a female that determine egg size are not clear. Although egg size often increases with female age (17 out of 37 studies), the change in egg size is generally less than 10%. Female mass and size rarely explain more than 20% of the variation in egg size within species. A female's egg size is not consistently related to other aspects of reproductive performance such as clutch size, laying date, or the pair's ability to rear young. Physiological characteristics of the female (e.g. endogenous protein stores, oviduct mass, rate of protein uptake by ovarian follicles) show more promise as potential determinants of egg size. With regards to the consequences of egg-size variation for offspring fitness, egg size is often correlated with offspring mass and size within the first week after hatching, but the evidence for more long-lasting effects on chick growth and survival is equivocal. In other oviparous vertebrates, the magnitude of egg-size variation within populations is often as great or greater than that observed within avian populations. Although there are much fewer estimates of the repeatability of egg size in other taxa, the available evidence suggests that egg size may be more flexible within individuals. Furthermore, in non-avian species (particularly fish and turtles), it is more common for female mass or size to explain a substantial proportion of the variation in egg size. Further research into the physiological basis of egg-size variation is needed to shed light on both the proximate and ultimate causes of intraspecific variation in this trait in birds.

Hatchery enhancement has been reported to result in an increase in egg size in coho salmon Oncorhynchus kisutch and a decline in egg size in Chinook salmon O. tshawytscha. Egg size may be directly influenced by selection, a larger egg size evolving as a consequence of hatchery incubation. Alternatively, a smaller egg size could evolve as a correlated response to fecundity selection, and a unidirectional change in egg size over time may reflect selection and an underlying genetic change in the population. To address this question, temporal trends in egg size were investigated for two hatchery‐enhanced populations of Chinook salmon from Vancouver Island, British Columbia. After the effect of female length variation was removed by standardizing egg sizes to a female of common length (the overall mean for each population), there was no temporal trend in egg size from the 1970s to 2008 for any of the hatchery‐enhanced populations evaluated. These results do not support a previous report of genetically based declines in egg size in hatchery‐enhanced Chinook salmon populations from this region.

Summary1. Temperature is considered one of the most important mediators of phenotypic plasticity in ectotherms, resulting in predictable changes in egg size. However, the fitness consequences of temperature‐induced plasticity in egg size are not well understood and are often assessed at mild temperatures, whereas in the field, extreme temperatures will occur.2. In this study we investigated egg size plasticity and fitness‐linked traits of eggs in response to temperature in the springtailOrchesella cincta. Eggs were oviposited at two temperatures (16 and 20 °C), and transplanted to 10, 15 and 20 °C for rearing, or exposed to temperature shock (−6 and 38 °C) to measure survival, development rate and growth.3. Eggs oviposited at 16 °C had a greater volume and dry weight than eggs oviposited at 20 °C but had similar lipid content. Eggs oviposited at 16 °C had high survival at all rearing temperatures, whereas survival of eggs oviposited at 20 °C declined at lower rearing temperatures and was significantly lower than survival of large eggs when reared at 10 and 15 °C. No difference in egg development rate was found between oviposition temperatures, although development rate increased with rearing temperature. Juveniles were 5–10% larger when hatching from eggs oviposited at 16 °C, independent of the temperature at which the eggs were reared.4. Eggs oviposited at 16 °C were more resistant to thermal stress, as their survival after exposure to cold or heat shock was higher than survival of eggs oviposited at 20 °C. However, egg development rate of the eggs that survived temperature shock was not different from the control, independent of egg oviposition temperature.5. Our results show that temperature‐induced plasticity inO. cinctaaffects fitness‐linked traits of offspring through increased hatchling size and increased thermal tolerance of eggs produced at lower temperatures.6. The increased resistance to thermal stress ofO. cinctaeggs oviposited at lower temperatures may provide a protective strategy in uncertain thermal environments.

According to the Climatic Variability Hypothesis [CVH], thermal plasticity should be wider in organisms from temperate environments, but is unlikely to occur in tropical latitudes where temperature fluctuations are narrow. In copepods, food availability has been suggested as the main driver of phenotypic variability in adult size if the range of temperature change is less than 14°C. Leptodiaptomus garciai is a calanoid copepod inhabiting Lake Alchichica, a monomictic, tropical lake in Mexico that experiences regular, narrow temperature fluctuations but wide changes in phytoplankton availability. We investigated whether the seasonal fluctuations of temperature and food produce phenotypic variation in the life-history traits of this tropical species. We sampled L. garciai throughout a year and measured female size, egg size and number, and hatching success, along with temperature and phytoplankton biomass. The amplitude of the plastic responses was estimated with the Phenotypic Plasticity Index. This index was also computed for a published dataset of 84 copepod populations to look if there is a relationship between the amplitude of the phenotypic plasticity of adult size and seasonal change in temperature. The temperature annual range in Lake Alchichica was 3.2°C, whereas phytoplankton abundance varied 17-fold. A strong pattern of thermal plasticity in egg size and adult female size followed the inverse relationship with temperature commonly observed in temperate environments, although its adaptive value was not demonstrated. Egg number, relative reproductive effort and number of nauplii per female were clearly plastic to food availability, allowing organisms to increase their fitness. When comparing copepod species from different latitudes, we found that the magnitude of thermal plasticity of adult size is not related to the range of temperature variation; furthermore, thermal plasticity exists even in environments of limited temperature variation, where the response is more intense compared to temperate populations.

Maternal effects are widespread and influence a variety of traits, for example, life history strategies, mate choice, and capacity to avoid predation. Therefore, maternal effects may also influence phenotypic plasticity of offspring, but few studies have addressed the relationship between maternal effects and phenotypic plasticity of offspring. We examined the relationship between a maternally influenced trait (egg size) and the phenotypic plasticity of the induction rate of the broad-headed morph in the salamander Hynobius retardatus. The relationship between egg size and the induction of the broad-headed morph was tested across experimental crowding conditions (densities of low conspecifics, high conspecifics, and high heterospecific anuran), using eggs and larvae from eight natural populations with different larval densities of conspecifics and heterospecifics. The broad-headed morph has a large mouth that enables it to consume either conspecifics or heterospecifics, and this ability gives survival advantages over the normal morph. We have determined that there is phenotypic plasticity in development, as shown by an increase in the frequency of broad-headed morph in response to an increase in the density of conspecifics and heterospecifics. This reaction norm differed between populations. We also determined that the frequency of the broad-headed morph is affected by egg size in which larger egg size resulted in expression of the broad-headed morph. Furthermore, we determined that selection acting on the propensity to develop the broad-headed morph has produced a change in egg size. Lastly, we found that an increase in egg size alters the reaction norm to favor development of the broad-headed morph. For example, an equal change in experimental density produces a greater change in the frequency of the broad-headed morph in larvae developing from large eggs than it does in larvae developing from small eggs. Population differences in plasticity might be the results of differences in egg size between populations, which is caused by the adaptive integration of the plasticity and egg size. Phenotypic plasticity can not evolve independently of maternal effects.

Fish species are known for their large phenotypic plasticity in life-history traits in relation to environmental characteristics. Plasticity allows species to increase their fitness in a given environment. Here we examined the life-history response of fish species after an abrupt change in their environment caused by the damming of rivers. Two reservoirs of different age, both situated on the Guiana Shield, were investigated: the young Petit-Saut Reservoir in French Guiana (14 years) and the much older Brokopondo Reservoir in Suriname (44 years). Six life-history traits in 14 fish species were studied and compared to their value in the Sinnamary River prior to the completion of Petit-Saut Reservoir. The traits analyzed were maximum length, absolute and relative length at first maturation, proportion of mature oocytes in ripe gonad, batch fecundity and mean size of mature oocytes. The results revealed a general increase of reproductive effort. All species showed a decrease in maximum length. Compared to the values observed before the dam constructions, eight species had larger oocytes and three species showed an increased batch fecundity. These observed changes suggest a trend towards a pioneer strategy. The changes observed in Petit-Saut Reservoir also seemed to apply to the 30 years older Brokopondo Reservoir suggesting that these reservoirs remain in a state of immaturity for a long time.

The Ability of Young and Old Hens to Change Shell Deposition with Sudden Natural Drastic Changes in Egg Size

Egg size affects larval size, growth rate, survival and fecundity with maternal fitness being maximized by a trade-off between egg size and fecundity. Production of a small number of large eggs maximizes female fitness under poor food conditions, as does a large number of small eggs under rich food conditions. Gnathopogon caerulescens (Honmoroko) spawns over a wide range of water temperature from spring to summer. Thus, we determined whether or not egg size varied with water temperature and how egg size influenced hatchling size at different water temperature. Changes in egg size strongly correlated with seasonal changes in water temperature around the lake, regardless of time and area. An experiment using eggs from the lake indicated that hatchling size has a significant positive relationship with egg size in water temperatures of 24 °C. On the other hand, a lower incubation temperature, similar to that likely to be encountered at the beginning of the spawning season, resulted in a smaller hatchling size, eggs requiring a longer time to hatch. At the beginning of the spawning season, therefore, egg size may have a lesser impact on hatching size in the natural environment of Honmoroko because of the relatively lower temperatures. Although fish larvae below a certain threshold of effective body size are generally not expected to have a high early survival rate due to, for example, a lower competitive ability or a high susceptibility to predation, seasonal egg size variation in Honmoroko may be adaptive so as to enhance the likehood of early survival by establishing an effective body size under changing water temperature conditions.

Reproductive schedules are a critical aspect of life history intrinsically linked to a species' ecology. We explored dynamic trajectories of daily fecundity, egg size, and egg fertility in three size classes of Hippodamia convergens Guerin-Meneville produced by varying larval access to food, eggs of Ephestia kuehniella Zeller. Adult pairs were held with ad libitum food and eggs were collected daily, counted, and a subsample measured. Egg fertility declined steeply over 25 clutches in small females, gradually in large females, but remained relatively stable in medium females. In small females, egg size and daily fecundity declined in a linear manner. There was no clear indication of an egg size-number tradeoff. In medium females, both egg size and daily fecundity peaked around the 16th day of oviposition, after which both declined. Large females began oviposition earlier and achieved peak egg size about day 7, and peak fecundity around day 12. Large females thus expressed a larger proportion of their reproductive effort early in adult life, a strategy inferred to be adaptive in the context of aphidophagy; a larger proportion of progeny would develop early in the exponential growth phase of the prey population when food is most abundant. Increases in egg size during this period may improve the survival of later-developing progeny; prey become scarce as aphid outbreaks decline and competition intensifies, favoring offspring with a larger body size at eclosion. Larval diet restriction appeared to constrain these presumably adaptive changes in egg size and daily fecundity, largely through effects on maternal body size.

Abstract. Measurements of egg number, egg size and total egg volume in relation to post-stripped weight, made on the eggs stripped from 12 separately maintained farm stocks of 4-year-old rainbow trout. Oncorhynchus mykiss Richardson, revealed highly significant differences between populations when analysed by regression and covariance statistical techniques. All of the stocks showed increases in fecundity with increasing fish size, with 61% (coefficient of determination or r2) of the variation in fecundity of the 12 stocks explained by the common regression on parental fish size. Comparison of the regressions of the different stocks revealed that the rate of increase of fecundity with increasing fish weight was constant throughout the weight range (i.e. the regressions had similar slopes) although there were highly significant differences in elevation, with some stocks producing almost twice as many eggs as some of the others. By contrast, the relationship of egg size with fish weight was much more variable with three of the stocks showing no significant regression of egg size on fish weight and a further two stocks with data poorly fitted to the regressions. Of the remaining seven stocks only one showed a significant difference in slope of the regression with modest increases in fish weight being associated with large increases in egg size. Only four of the remaining stocks of fish showed significant differences in the elevation of their regressions, i.e. the egg sizes were significantly different after the potentially conflicting influence of fish size was partitioned using covariance techniques. Assessments of total egg volume produced far improved regressions on post-strip weight with 75% (r2) of the variation in the pooled data of the 12 stocks being described by the line of the common regression. In general, there appeared to be a “trade-off” between egg number and size with the most and least fecund fish producing somewhat smaller and larger eggs respectively. All 12 regressions of totai egg volume on fish weight showed a common slope. However, there were significant differences in elevation of the regressions, with some of the stocks of trout producing up to a 55% greater volume of eggs. There appeared to be no relationship between the differences in egg number, size or total egg volume and the type or geograpfiical location of the farms on which the stocks were held. Although it remains to be determined to what extent these differences in reproductive performance are Inheritable, it is recommended that these characteristics should be taken into account by hatcheries in tbe design of long-term broodstock selection programmes for rainbow trout.

Larvae of two species of sea urchins (Strongylocentrotus droebachiensis and S. purpuratus) differ in initial form and in the rate of development. To determine whether these differences are attributable to the large interspecific difference in egg size, we experimentally reduced egg size by isolating blastomeres from embryos. The rate of development of feeding larvae derived from isolated blastomeres was quantified using a novel morphometric method. If the differences early in the life histories of these two species are due strictly to differences in egg size, then experimental reduction of the size of S. droebachiensis eggs should yield an initial larval form and rate of development similar to that of S. purpuratus. Our experimental manipulations of egg size produced three clear results: 1) smaller eggs yielded larvae that were smaller and had simpler body forms, 2) smaller eggs resulted in slower development through the early feeding larval stages, and 3) effects of egg size were restricted to early larval stages. Larvae from experimentally reduced eggs of the larger species had rates of development similar to those of the smaller species. Thus, cytoplasmic volumes of the eggs, not genetic differences expressed during development, account for differences in larval form and the rate of form change. This is the first definitive demonstration of the causal relationship between egg size (parental investment per offspring) and life-history characteristics in marine benthic invertebrates. Because larval form influences feeding capability, the epigenetic effects of egg size on larval form are likely to have important functional consequences. Adaptive evolution of egg size may be constrained by the developmental relationships between egg size and larval form: evolutionary changes in egg size alone can result in concerted changes in larval form and function; likewise evolutionary changes in larval form and function can be achieved through changes in egg size. These findings may have broader implications for other taxa in which larval morphology and, consequently, performance may be influenced by changes in egg size.

Although the temperature‐size rule, that is, an increase in egg (and body) size at lower temperatures, applies almost universally to ectotherms, the developmental mechanisms underlying this consistent pattern of phenotypic plasticity are hitherto unknown. By investigating ovarian dynamics and reproductive output in the tropical butterfly Bicyclus anynana (Butler) (Lepidoptera: Nymphalidae: Satyrinae) in relation to oviposition temperature and mating status, we tested the relevance of several competing hypotheses for temperature‐mediated variation in egg size and number. As expected, females ovipositing at a lower temperature laid fewer but larger eggs than those ovipositing at a higher temperature. Despite pronounced differences in egg‐laying rates, oocyte numbers were equal across temperatures at any given time, while oocyte size increased at the lower temperature. In contrast, there were greatly reduced oocyte numbers in mated compared to virgin females. Our results indicated that temperature‐mediated plasticity in egg size cannot be explained by reduced costs of somatic maintenance at lower temperatures, enabling the allocation of more resources to reproduction (reproductive investment was higher at the higher temperature). Furthermore, there was no indication for delayed oviposition (no accumulation of oocytes at the lower temperature, in contrast to virgin females). Rather, low temperatures greatly reduced the oocyte production (i.e., differentiation) rate and prolonged egg‐maturation time, causing low egg‐laying rates. Our data thus suggested that oocyte growth is less sensitive to temperature than oocyte production, resulting in a lower number of larger eggs at lower temperatures.

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.

Temperature is considered one of the most important mediators of phenotypic plasticity in ectotherms. However, the costs and benefits shaping the evolution of different thermal responses are poorly elucidated. One of the possible constraints to phenotypic plasticity is its intrinsic genetic cost, such as genetic linkage or pleiotropy. Genetic coupling of the thermal response curves for different life history traits may significantly affect the evolution of thermal sensitivity in thermally fluctuating environments. We used the collembolan Orchesella cincta to study if there is genetic variation in temperature-induced phenotypic plasticity in life history traits, and if the degree of temperature-induced plasticity is correlated across traits. Egg development rate, juvenile growth rate and egg size of 19 inbred isofemale lines were measured at two temperatures. Our results show that temperature was a highly significant factor for all three traits. Egg development rate and juvenile growth rate increased with increasing temperature, while egg size decreased. Line by temperature interaction was significant for all traits tested; indicating that genetic variation for temperature-induced plasticity existed. The degree of plasticity was significantly positively correlated between egg development rate and growth rate, but plasticity in egg size was not correlated to the other two plasticity traits. The findings suggest that the thermal plasticities of egg development rate and growth rate are partly under the control of the same genes or genetic regions. Hence, evolution of the thermal plasticity of traits cannot be understood in isolation of the response of other traits. If traits have similar and additive effects on fitness, genetic coupling between these traits may well facilitate the evolution of optimal phenotypes. However, for this we need to know the selective forces under field conditions.

This study examined 23 years of egg size data from a population entirely made up of hatchery‐maintained fall Chinook Salmon Oncorhynchus tshawytschain Lake Oahe, South Dakota. Egg size data, obtained by water‐displacement during hatchery egg inventories, ranged from 4.2 to 6.6 eggs/mL over the 23 years. However, linear regression of data from either pooled lots of eggs (y = 0.0035x – 1.8116; P = 0.727) or individual spawns (y = 0.0003x + 5.3037; P = 0.907) indicated no significant change in mean egg size over time. The lack of change in egg size of Lake Oahe fall Chinook Salmon over the 23 years of this study indicates there was no hatchery‐induced evolutionary impacts on egg size.