Size-number trade-off and optimal offspring size for offspring produced sequentially using a fixed amount of reserves

We analysed the nature of size-number trade-off of offspring when multiple cohorts of such offspring are produced sequentially using a fixed amount of reserves. In the model, we incorporated sink-limitation in the resource absorption rate of offspring from the mother tissue and the loss of resources by maintenance respiration. We found that the later the initiation of a cohort, the greater the cost of producing a cohort with the same size and number of offspring. This is due to the loss of resources by maintenance respiration during the period from the beginning of reproduction to the initiation of the cohort. Also, the extra cost increases with an increase in the specific maintenance respiration rate. Thus, resources lost to respiration over time reduces the fitness value of producing late cohorts. Hence, it is advantageous to produce all offspring simultaneously unless there are fitness advantages of producing offspring sequential which overcome this cost or constraints preventing simultaneous production. Sequentially offspring production evolves if there is a constraint on the number of offspring of each cohort. With this constraint, the optimal offspring size decreases with the production sequence of cohorts.     

Optimal offspring sizes in small litters

Summary Numerous evolutionary models explore the trade-off between offspring size and offspring number. However, such models often fail when the number of offspring is small because optimal litter size (or litter size at optimal offspring size) may fall between the necessarily integer values for real litters. This paper extends a classic model for optimal investment per offspring to the case of small litters and predicts that range in offspring size and the largest (smallest) offspring size should decline (increase) with increased litter size. Application of the model to egg size data from a poeciliid fish,Gambusia hubbsi, reveals a surprisingly close approximation to the largest offspring size and variation in offspring size at small litter sizes.     

The influence of a positive correlation between clutch size and offspring fitness on the optimal offspring size

Summary The effect is modeled of a positive relationship between clutch size and offspring fitness on the optimal investment in offspring. In species which meet the assumptions of the model, the model predicts a positive correlation between maternal resource level and offspring size. If larger mothers are able to allocate more resources to offspring, then the model would also predict a positive correlation between maternal size and offspring size when the assumptions of the model are met. Thus, this model may help explain both among and within individual variation in offspring size. When offspring are produced in groups and the number of offspring killed per clutch is limited by predator satiation, offspring in larger clutches may experience a higher probability of survival. Such a life style may be found in animals such as sea turtles. Offspring size is positively correlated with maternal size in some members of this group.     

Clonal plasticity in response to nutrient availability in the pseudoannual herb,Trientalis europaea L.

Trientalis europaea plants were subjected to four levels of nutrient availability to test two hypotheses: (1) Clonal morphology of pseudoannuals with short-lived rhizomes is responsive to nutrient availability in terms of spacer length (a negative correlation expected) and branching intensity (a positive correlation expected), and (2) the size of the hibernacles of T. europaea shows a positive correlation with nutrient availability. The results support the first hypothesis, since the primary spacers of T. europaea were longer at lower nutrient levels while the branching intensity of the primary rhizomes increased at increasing nutrient supply. The second hypothesis was not confirmed, however; the plants produced fewer, but bigger hibernacles at lower levels of nutrient availability. The ecological significance of the results is discussed in the context of foraging for essential resources and habitat-related effects of hibernacle size on survival and establishment chances of daughter ramets.     

Differences in allocation patterns in clonal and sexual offspring in a woodland pseudo-annual

We compared the patterns of allocation to reproduction among seed-derived and clonal offspring of a woodland pseudo-annual. Pseudo-annuals are clonal plants which survive the winter only as seeds and hibernacles produced by the rhizome system. Previous studies indicate that flowering is related to the size of these hibernacles. Since seedlings do not have a hibernacle, we did not expect that these plants would reproduce sexually. Assuming a trade-off between sexual and asexual reproduction, and assuming a linear relationship between vegetative plant weight and weight of all reproductive structures (i.e., rhizomes, hibernacles, inflorescences, and seeds), we expected that seed-derived plants would have a stronger biomass allocation to rhizomes and hibernacles. Since resource supply affects plant size, and thus hibernacle and seed production, we also subjected the plants to different levels of shade. At the start of the experiment seed-derived and clonal offspring hardly differed in total fresh weight. At the final harvest in September seed-derived and clonal offspring did not differ in vegetative plant weight (i.e., leaves, stems, and roots). Only light availability significantly affected these plant structures. As predicted, seed-derived plants did not flower in either of the light treatments. Seed-derived plants allocated more biomass to rhizomes and hibernacles, but this was only significant in the highest-light treatment. This result was due only to an increase in the number of hibernacles. Dry weight of single hibernacles was not affected by plant type. The ecological implications of this allocation pattern are discussed. Received: 2 October 1997 / Accepted: 8 March 1998     

Offspring size-number strategy in the bethylid parasitoid Laelius pedatus

I observed clutch size and body size of resulting offspringfor the parasitoid Laelies pedatus (Say) (Hymenoptera: Bethylidae)on hosts of different sizes. Results were compared with thepredictions of offspring size-number models and clutch-sizemodels. Larger clutches were laid on larger hosts. However,even after females had adjusted dutch size to the size of thehost, offspring size was larger in larger broods. The variancein offspring size between broods decreased with increasing dutchsize as expected, but the decrease was smaller than predictedby Charaov and Downhower's trade-off invariant rule. Theorypredicts such trends when the shape of the trade-off betweenper capita investment and per capita offspring fitness dependson dutch size or host size. By observing how this assumptionmight apply to bethylid wasps, I generate a number of testablehypotheses to explain the observed trends.     

Experimentally reducing clutch size reveals a fixed upper limit to egg size in snakes, evidence from the king ratsnake, Elaphe carinata

Snakes are free of the pelvic girdle's constraint on maximum offspring size, and therefore present an opportunity to investigate the upper limit to offspring size without the limit imposed by the pelvic girdle dimension. We used the king ratsnake (Elaphe carinata) as a model animal to examine whether follicle ablation may result in enlargement of egg size in snakes and, if so, whether there is a fixed upper limit to egg size. Females with small sized yolking follicles were assigned to three manipulated, one sham-manipulated and one control treatments in mid-May, and two, four or six yolking follicles in the manipulated females were then ablated. Females undergoing follicle ablation produced fewer, but larger as well as more elongated, eggs than control females primarily by increasing egg length. This finding suggests that follicle ablation may result in enlargement of egg size in E. carinata. Mean values for egg width remained almost unchanged across the five treatments, suggesting that egg width is more likely to be shaped by the morphological feature of the oviduct. Clutch mass dropped dramatically in four- and six-follicle ablated females. The function describing the relationship between size and number of eggs reveals that egg size increases with decreasing clutch size at an ever-decreasing rate, with the tangent slope of the function for the six-follicle ablation treatment being -0.04. According to the function describing instantaneous variation in tangent slope, the maximum value of tangent slope should converge towards zero. This result provides evidence that there is a fixed upper limit to egg size in E. carinata.     

Factors influencing offspring traits in the oviparous multi-clutched lizard,<Emphasis Type=Italic>Calotes versicolor</Emphasis> (Agamidae)

The determinants of offspring size and number in the tropical oviparous multi-clutched lizard,Calotes versicolor, were examined using both univariate and multivariate (path) analyses. InC. versicolor maternal snout-vent length (SVL) and body condition influence clutch mass and clutch size but have no significant influence on offspring size. The positive effect of maternal SVL and body condition on offspring number is counterbalanced by a negative effect of breeding time on egg mass. In fact, breeding time directly influences the offspring body mass and condition through variation in the egg mass. There is a trade-off between offspring mass and condition with offspring number, and breeding time influences both. Offspring hatched from the eggs of early (May–June) or mid (July–August) breeding periods invariably show lower mass and condition than those hatched from the eggs of late breeding season (September–October). Yet, there is no variation in offspring SVL among early, mid and late clutches. Thus, inC. versicolor offspring SVL is optimized while body mass and condition are not optimized.     

Does follicle excision always result in enlargement of offspring size in lizards?

An experimental reduction of offspring number has been reported to result in enlargement of offspring size in lizards. We applied the “follicle excision” technique to a lacertid lizard (Takydromus septentrionalis) to examine whether this effect is generalisable to lizards. Of the 82 females that produced 3 successive clutches in the laboratory, 23 females underwent follicle excision after they oviposited the first clutch. Follicle excision reduced clutch size, but did not alter egg size. This result indicates that egg size is not altered during vitellogenesis in T. septentrionalis. Females undergoing follicle excision produced a third clutch (a second post-surgical clutch) as normally as did control females. Females switched from producing more but smaller eggs early in the breeding season to fewer but larger eggs later in the season. Our results indicate that female T. septentrionalis maximize reproductive success by diverting an optimal, rather than a higher, fraction of the available energy to individual offspring. This optimized allocation of the available energy to offspring production explains why follicle excision does not result in enlargement of egg size in this species. Our study provides evidence that an experimental reduction of offspring number does not always result in enlargement of offspring size in lizards.     

Energetics of reproduction: consequences of divergent selection on egg size, food limitation, and female age for egg composition and reproductive effort in a butterfly

Using lines artificially selected on egg size and being subjected to a restricted and an unrestricted feeding treatment, we examined the relationships between egg size, egg number, egg composition, and reproductive investment in the butterfly Bicyclus anynana . Despite a successful manipulation of egg size, correlated responses to selection in larval time, pupal mass, pupal time, longevity, fecundity, or the amount of energy allocated to reproduction were virtually absent. Thus, there was no indication for an evolutionary link between offspring size and reproductive investment. Egg composition, in contrast, was affected by selection, with larger eggs containing relatively more lipid and water, but less protein and energy compared to smaller eggs. Hence, females producing large eggs did not have to sacrifice fecundity due to adjustments in egg composition. Food limitation per se caused only minor changes in egg composition, and there was no general reduction in egg provisioning with female age. The latter was restricted to food-limited females, whereas egg quality remained remarkably similar throughout the females' life in control groups. We conclude that neglecting changes in biochemical egg composition, depending on genetic background, food availability, and female age, may introduce substantial error when estimating reproductive effort, and may ultimately lead to invalid conclusions.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 403–418.     

密点麻蜥的两性异形和雌性繁殖

蜥蜴繁殖成功率与其形态特征有密切的关系。作者在内蒙古乌拉特后旗采集密点麻蜥(Eremias multio-cellata) ,定量研究该种形态特征的两性异形和雌体繁殖特征,检验与成体形态特征相关的两性繁殖成功率差异是否能促进两性异形的进化。密点麻蜥成体个体大小无显著的两性差异,但头部大小两性差异显著;雄性个体的头长和头宽均大于体长相同的雌性成体。繁殖雌体于五、六月份排卵;在实验室条件下,雌体在六月下旬至七月下旬之间产仔。该种雌体年产单窝仔,每窝2 -4仔。窝仔重与雌体体长呈正相关,但雌体体长仅能解释很少一部分(约19 %)窝仔重的变异。窝仔数和幼仔重均与雌体体长无关。幼仔重与相对生育力(相对于雌体体长的窝仔数)呈显著的负相关,表明该种蜥蜴存在后代数量-大小之间的权衡。密点麻蜥雄体和雌体向较大体型方向进化的选择压力均相对较弱,与成体头部大小相关的两性繁殖成功率的差异是导致该种蜥蜴头部大小两性异形进化的主要原因[动物学报52 (2) : 250 -255 , 2006]。     

The evolution of offspring size and number: a test of the Smith-Fretwell model in three species of crickets

Most models of parental investment in offspring assume a trade-off between propagule size and number, and an increasing concave down function relating offspring fitness to propagule size. In this study, we test these two fundamental assumptions, using three closely related species of crickets, Gryllus firmus, G. veletis, and G. pennsylvanicus. Egg weight, 35-day fecundity and 35-day egg biomass were estimated in a population of each species, and the relationships between these reproductive traits and date of egg laying and body size were estimated. The relationships between egg weight and offspring survival were also sought for eggs buried at different depths, soil moistures, and soil types (G. firmus and G. veletis), as well as in the field (G. pennsylvanicus). A trade-off between egg weight and 35-day fecundity was revealed in a multivariate analysis taking into account among-species variation in egg weight and body size. Independent of the environmental conditions affecting the eggs, a positive correlation existed between the number of larvae that emerged from the soil and propagule weight in each species. Therefore, these findings provide partial support for the assumptions considered in the models mentioned above. A single optimal egg size was favored in two out of the three sets of conditions in which the functions relating egg weight to larval survival could be derived. The conditions encountered by the eggs, however, influenced the average survival of the larvae, as well as the shape of the relationship between egg weight and offspring survival. This suggests that cricket eggs frequently face heterogeneous environments with respect to egg and hatchling survival; the implication of habitat heterogeneity on the evolution of an optimal egg size is considered. The relationships between the reproductive components and female age and size, as well as between egg size and variation in cricket life-history, are discussed in an ecological and evolutionary context.     

Meta-ecosystems and biological energy transport from ocean to coast: the ecological importance of herring migration

Ecosystems are not closed, but receive resource subsidies from other ecosystems. Energy, material and organisms are moved between systems by physical vectors, but migrating animals also transport resources between systems. We report on large scale energy transport from ocean to coast by a migrating fish population, the Norwegian spring-spawning (NSS) herring Clupea harengus. We observe a rapid body mass increase during parts of the annual, oceanic feeding migration and we use a bioenergetics model to quantify energy consumption. The model predicts strong seasonal variation in food consumption with a marked peak in late May to July. The copepod Calanus finmarchicus is the most important prey and 23×10⁶ tonnes (wet weight) of C. finmarchicus is consumed annually. The annual consumption-biomass ratio is 5.2. During the feeding migration 17% of consumed energy is converted to body mass. The biomass transported to the coast and left as reproductive output is estimated from gonad weight and is about 1.3×10⁶ tonnes for the current population. This transport is to our knowledge the world’s largest flux of energy caused by a single population. We demonstrate marked temporal variation in transport during the last century and discuss the effects of NSS herring in the ocean, as a major consumer, and at the coast, where eggs and larvae are important for coastal predators. In particular, we suggest that the rapid decline of lobster Homarus gammarus landings in Western Norway during the 1960s was related to the collapse of NSS herring. We also discuss spatial variation in energy transport caused by changed migration patterns. Both climate and fisheries probably triggered historical changes in the migration patterns of NSS herring. New migration routes emerge at the level of individuals, which in turn determines where resources are gathered and delivered, and therefore, how meta-ecosystems function.