Constructional morphology of strombid gastropods

The extreme diversity in shell shape of strombid gastropods is interpreted as the result of three independent factors: (1) The terminal growth pattern of the Strombidae allows the circumvention of geometric constraints on shell morphology found in gastropods with continuous or periodic growth patterns. (2) Shell morphology in the Strombidac is adaptive in epifaunal locomotion, burrowing. infaunal or semi-infaunal habits, and passive protection from predators. Specialization for one of these functions often conflicted with the others. thus bringing about a forced 'choice' among mutually exclusive morphological characters. (3) Conservatism in life habits and anatomy of the soft parts has allowed the multiple evolution of extreme shell morphologies, as well as the secondary return to relativcly unspecialized morphologies. □ Constructional morphology, functional morphology. growth. behaviour. evolution, locomotion, burrowing, predation, exoskeleton. shell. Mollusca. Gastropoda. Strombacea. Strombidae.     

Differential growth rates and calcium-allocation strategies in the garden snail Cantareus aspersus

An optimal division of a key resource between growth and reproductionis expected to produce consistent life history schedules inhabitats where its supply is highly predictable. However, differentialgrowth rates are found between populations and within broodsof Cantareus aspersus, a simultaneous hermaphrodite for whichthe reproductive benefits of a large body size may favour rapidgrowth. Although energy is usually assumed to be the limitingresource in allocation theory, calcium limits the distribution,growth and reproduction of snails. This is a very consistentresource and populations may have allocation strategies whichreflect availability in their habitats. Three experiments comparedCa allocation in the progeny of six populations from Ca-richand Ca-poor habitats. In the first, 100 d-old juveniles werecompared between populations for their shell/soft-tissue dryweight ratio, their allocation of Ca to each compartment, andthe variability within broods. The second measured growth, foodconsumption and shell ratios in growth trials of three populationson low Ca. Thirdly, five populations were compared on abundantor excess Ca. The relationship of shell Ca with soft-tissuelevels differs between populations, but shell ratios changedwith Ca availability in all populations. Most favoured soft-tissuegrowth when dietary Ca is low, but one population (LE) alwayshad the highest shell ratios in these trials. Ca in the parentalhabitat was not a good predictor of juvenile-allocation strategies,but the consistency of LE shell ratios across several broodssuggests theirs may be an inherited trait. LE has faster growthrates and a preference for shell building, which probably representsa strategy for early reproduction. The robustness of a snail'sshell may thus be more indicative of its reproductive strategyrather than Ca availability in its habitat. (Received 18 May 2006; accepted 21 December 2006)     

Dynamic state-dependent modelling predicts optimal usage patterns of responsive defences

Chemical defences against predation often involve responses to specific predation events where the prey expels fluids, such as haemolymph or gut contents, which are aversive to the predator. The common link is that each predation attempt that is averted results in an energetic cost and a reduction in the chemical defences of the prey, which might leave the prey vulnerable if the next predation attempt occurs soon afterwards. Since prey appear to be able to control the magnitude of their responses, we should expect them to trade-off the need to repel the current threat against the need to preserve defences against future threats and conserve energy for other essential activities. Here we use dynamic state-dependent models to predict optimal strategies of defence deployment in the juvenile stage of an animal that has to survive to maturation. We explore the importance of resource level, predator density, and the costs of making defences on the magnitude of the responses and optimal age and size at maturation. We predict the patterns of investment and the magnitude of the deployment of defences to potentially multiple attacks over the juvenile period, and show that responses should be smaller when the costs of defences and/or predation risk are higher. The model enables us to predict that animals in which defences benefit the adult stage will employ different strategies than those that do not use the same defences as adults, and thereby experience a smaller reduction in body size as a result of repeated attacks. We also explore the effect of the importance of adult size, and find that the sex and mating system of the prey should also affect defensive strategies. Our work provides the first predictive theory of the adaptive use of responsive defences across taxa.     

Sexual size dimorphism in species with asymptotic growth after maturity

If animals mature at small sizes and then grow to larger asymptotic sizes, many factors can affect male and female size distributions. Standard growth equations can be used to study the processes affecting sexual size dimorphism in species with asymptotic growth after maturity. This paper first outlines the effects of sex differences in growth and maturation patterns on the direction and degree of sexual dimorphism. The next section considers the effects of variation in age structure or growth rates on adult body sizes and sexual size dimorphism. Field data from a crustacean, fish, lizard and mammal show how information on a species' growth and maturation patterns can be used to predict the relationships between male size, female size and sexual size dimorphism expected if a series of samples from the same population simply differed with respect to their ages or growth rates. The last section considers ecological or behavioural factors with different effects on the growth, maturation, survival or movement patterns of the two sexes. This study supports earlier suggestions that information on growth and maturation patterns may be useful, if not essential, for comparative studies of sexual size dimorphism in taxa with asymptotic growth after maturity.     

Optimal patterns of growth and reproduction for perennial plants with persisting or not persisting vegetative parts

Summary Optimal allocation of energy to growth, reproduction and storage was considered for perennial plants differing in the proportion of vegetative structures persisting over winter and/or in the amount of resources which can be relocated to storage before abscission of some organs. It was found that for every mortality level there exists a critical proportion of persistent organs. Below this critical value it is optimal to grow without reproduction for the first years until a characteristic size is reached; afterwards, that size is maintained year after year and all extra resources are devoted to reproduction. Some storage is also necessary to maintain constant size. If the proportion of retained vegetative mass is above the critical value, the optimal strategy is gradual growth to an asymptotic size, with growth and reproduction occurring in several years following maturation. In this case real storage occurs only until maturation is reached, then storage is realized only by energy relocation from the vegetative body. Although the optimal solution changes abruptly qualitatively at a given proportion of resources saved from year to year, further growth of this proportion above the critical level brings about a greater difference between size reached at maturity and final size. The predictions of the model seem to follow the pattern of nature qualitatively.     

Unexpected discontinuities in life-history evolution under size-dependent mortality

In many organisms survival depends on body size. We investigate the implications of size-selective mortality on life-history evolution by introducing and analysing a new and particularly flexible life-history model with the following key features: the lengths of growth and reproductive periods in successive reproductive cycles can vary evolutionarily, the model does not constrain evolution to patterns of either determinate or indeterminate growth, and lifetime number and sizes of broods are the outcomes of evolutionarily optimal life-history decisions. We find that small changes in environmental conditions can lead to abrupt transitions in optimal life histories when size-dependent mortality is sufficiently strong. Such discontinuous switching results from antagonistic selection pressures and occurs between strategies of early maturation with short reproductive periods and late maturation with long reproductive cycles. When mortality is size-selective and the size-independent component is not too high, selection favours prolonged juvenile growth, thereby allowing individuals to reach a mortality refuge at large body size before the onset of reproduction. When either component of mortality is then increased, the mortality refuge first becomes unattractive and eventually closes up altogether, resulting in short juvenile growth and frequent reproduction. Our results suggest a new mechanism for the evolution of life-history dimorphisms.     

Optimal defense strategy against herbivory in plants: Conditions selecting for induced defense, constitutive defense, and no-defense

To examine the conditions selecting for induced defense, constitutive defense, and no-defense, we developed a model of plant defense strategy against herbivory. In the model, a plant consists of two modules between which signal inducing defense compounds can be translocated. We assume three strategies: plants produce defense compounds responding to herbivory (induced defense), they have the compounds beforehand (constitutive defense), and they never produce the compounds (no-defense). We found that no-defense is optimal if the amount of biomass lost due to herbivory is small because of the growth cost of having defense compounds. The constitutive defense is optimal if the amount of biomass lost is not so small and the probability of herbivory is high. If the biomass loss is not so small but the probability of herbivory is low, the induced defense or no-defense is optimal. When the induced defense is optimal, the probability of herbivory necessarily increases in plants once herbivory has occurred. If the probability stays the same, no-defense is optimal. Thus, the behavior of herbivores, i.e., whether they remain around a plant and attack it repeatedly, affects the evolution of the induced defense.     

Development of natural growth regimes for hatchery-reared steelhead to reduce residualism, fitness loss, and negative ecological interactions

Wild steelhead (Oncorhynchus mykiss) typically spend two or more years in freshwater before migrating to sea, but hatchery steelhead are almost ubiquitously released as yearlings. Their large size at release coupled with life history pathways that include both male and female maturation in freshwater present ecological risks different from those posed by hatchery populations of Pacific salmon. Yearling hatchery reared steelhead that fail to attain minimum thresholds for smoltification or exceed thresholds for male maturation tend to ‘residualize’ (i.e., remain in freshwater). Residuals pose ecological risks including size-biased interference competition and predation on juvenile salmon and trout. Three hatchery populations of steelhead in Hood Canal, WA were reared under growth regimes designed to produce a more natural age at smoltification (age-2) to aid in rebuilding their respective natural populations. Mean smolt sizes and size variability at age-2 were within the range of wild smolts for two of the three populations. The third population reared at a different facility under similar temperatures exhibited high growth rate variability and high male maturation rates (20% of all released fish). Experimentally comparing age-1 and age-2 smolt programs will help identify optimal rearing strategies to reduce the genetic risk of domestication selection and reduce residualism rates and associated negative ecological effects on natural populations. Investigations of Winthrop National Fish Hatchery summer-run steelhead will measure a) selection on correlated behavioral traits (‘behavioral syndromes’), b) degree of smoltification, c) changes in hormones that regulate gonad growth at key developmental stages, and d) conduct extensive post-release monitoring of fish reared under each growth regime.     

Optimizing time and resource allocation trade-offs for investment into morphological and behavioral defense

Prey organisms are confronted with time and resource allocation trade-offs. Time allocation trade-offs partition time, for example, between foraging effort to acquire resources and behavioral defense. Resource allocation trade-offs partition the acquired resources between multiple traits, such as growth or morphological defense. We develop a mathematical model for prey organisms that comprise time and resource allocation trade-offs for multiple defense traits. Fitness is determined by growth and survival during ontogeny. We determine optimal defense strategies for environments that differ in their resource abundance, predation risk, and defense effectiveness. We compare the results with results of simplified models where single defense traits are optimized. Our results indicate that selection acts in favor of integrated traits. The selective advantage of expressing multiple defense traits is most pronounced at intermediate environmental conditions. Optimizing single traits generally leads to a more pronounced response of the defense traits, which implies that studying single traits leads to an overestimation of their response to predation. Behavioral defense and morphological defense compensate for and augment each other depending on predator densities and the effectiveness of the defense mechanisms. In the presence of time constraints, the model shows peak investment into morphological and behavioral defense at intermediate resource levels.     

Effect of growth rate and hydrophobicity on bacteria surviving protozoan grazing

Measurements were made of the predation by Tetrahymena thermophila on several bacterial species in media containing heat-killed Escherichia coli cells to serve as an alternative prey. If grazing pressure was initially not intense on a mixture of bacterial species, the species that survived protozoan feeding at greater densities were those that grew quickly before the onset of active predation. If members of several species were incubated individually at similar initial densities with actively grazing T. thermophila, some species survived at ca. 10(4)/ml, some survived at ca. 10(2)/ml, and others were eliminated. Members of the first two groups but not the third group were able to multiply in the medium in the absence of the protozoan, but the growth rates in the protozoan-free medium did not correlate with the number of survivors. However, the species that persisted at the higher densities possessed highly hydrophobic cell surfaces. The size of the surviving population of four bacterial species whose growth was prevented by chloramphenicol correlated with the initial cell density that was incubated with T. thermophila. It is concluded that the individual species surviving predation on a mixture of species is related to the capacity of the bacterium to grow, the hydrophobicity of its cell surface, and the population density of the species before the onset of intense grazing.     

A mathematical consideration for the optimal shell change of hermit crab

Shell of the adult hermit crab has some important roles for its fitness. In the same time, the shell size often limits the body growth of its owner. To grow the body size larger, the individual must change the shell to another larger shell. If the individual cannot get another larger one, the individual has to suppress the body size growth as the occupied shell size allows. Growth suppression would result in the lower fitness. With a simple mathematical model, we consider the criterion about whether the individual should try to change the shell or not in order to get the higher fitness. We show that the optimality of a shell change behavior has a relation with the body size and the season length for the shell change. They also affect the optimal timing for the shell change. It is implied that the probability of the success in a shell change and the cost for the shell change behavior do not affect the optimal timing for the shell change at all but significantly do the optimality of the behavioral choice.     

Effect of growth rate and hydrophobicity on bacteria surviving protozoan grazing.

Measurements were made of the predation by Tetrahymena thermophila on several bacterial species in media containing heat-killed Escherichia coli cells to serve as an alternative prey. If grazing pressure was initially not intense on a mixture of bacterial species, the species that survived protozoan feeding at greater densities were those that grew quickly before the onset of active predation. If members of several species were incubated individually at similar initial densities with actively grazing T. thermophila, some species survived at ca. 10(4)/ml, some survived at ca. 10(2)/ml, and others were eliminated. Members of the first two groups but not the third group were able to multiply in the medium in the absence of the protozoan, but the growth rates in the protozoan-free medium did not correlate with the number of survivors. However, the species that persisted at the higher densities possessed highly hydrophobic cell surfaces. The size of the surviving population of four bacterial species whose growth was prevented by chloramphenicol correlated with the initial cell density that was incubated with T. thermophila. It is concluded that the individual species surviving predation on a mixture of species is related to the capacity of the bacterium to grow, the hydrophobicity of its cell surface, and the population density of the species before the onset of intense grazing.     

Impact of environmental covariation in growth and mortality on evolving maturation reaction norms

Maturation age and size have important fitness consequences through their effects on survival probabilities and body sizes. The evolution of maturation reaction norms in response to environmental covariation in growth and mortality is therefore a key subject of life-history theory. The eco-evolutionary model we present and analyze here incorporates critical features that earlier studies of evolving maturation reaction norms have often neglected: the trade-off between growth and reproduction, source-sink population structure, and population regulation through density-dependent growth and fecundity. We report the following findings. First, the evolutionarily optimal age at maturation can be decomposed into the sum of a density-dependent and a density-independent component. These components measure, respectively, the hypothetical negative age at which an individual's length would be 0 and the delay in maturation relative to this offset. Second, along any growth trajectory, individuals mature earlier when mortality is higher. This allows us to deduce, third, how the shapes of evolutionarily optimal maturation reaction norms depend on the covariation between growth and mortality (positive or negative, linear or curvilinear, and deterministic or probabilistic). Providing eco-evolutionary explanations for many alternative reaction-norm shapes, our results appear to be in good agreement with current empirical knowledge on maturation dynamics.     

Environmental influences on the evolution of growth and developmental rates in passerines

Abstract.— The reasons why growth and developmental rates vary widely among species have remained unclear. Previous examinations of possible environmental influences on growth rates of birds yielded few correlations, leading to suggestions that young may be growing at maximum rates allowed within physiological constraints. However, estimations of growth rates can be confounded by variation in relative developmental stage at fledging. Here, we re-estimate growth rates to control for developmental stage. We used these data to examine the potential covariation of growth and development with environmental variation across a sample of 115 North American passerines. Contrary to previous results, we found that growth rates of altricial nestlings were strongly positively correlated to daily nest predation rates, even after controlling for adult body mass and phylogeny. In addition, nestlings of species under stronger predation pressure remained in the nest for a shorter period, and they left the nest at lower body mass relative to adult body mass. Thus, nestlings both grew faster and left the nest at an earlier developmental stage in species with higher risk of predation. Growth patterns were also related to food, clutch size, and latitude. These results support a view that growth and developmental rates of altricial nestlings are strongly influenced by the environmental conditions experienced by species, and they generally lend support to an adaptive view of interspecific variation in growth and developmental rates.     

Target size and optimal life history when individual growth and energy budget are stochastic

I extend my previous work on life history optimization when body mass is divided into reserves and structure components. Two important innovations are: (1) effect of finite target size on optimal structural growth; (2) incorporating reproduction in the optimization objective. I derive optimal growth trajectories and life histories, given that the individual is subject to both starvation mortality and exogenous hazards (e.g., predation). Because of overhead costs in building structural mass, it is optimal to stop structural growth close to the target size, and to proceed only by accumulating reserves. Higher overhead costs cause earlier cessation of structural growth and smaller final structures. Semelparous reproduction also promotes early cessation of structural growth, compared to when only survival to target size is maximized. In contrast, iteroparous reproduction can prolong structural growth, resulting in larger final structures than in either the survival or the semelparous scenarios. Increasing the noise in individual growth lowers final structural mass at small target sizes, but the effect is reversed for large target sizes. My results provide predictions for comparative studies. I outline important consequences of my results to additional important evolutionary questions: evolution of sexual dimorphism, optimization of clutch size and evolution of progeny and adult sizes.     

The growth pattern of chimpanzees: Somatic growth and reproductive maturation inPan troglodytes

Growth of chimpanzees reared at the Kumamoto Primates Park of Sanwa Kagaku Kenkyusho Co. Ltd. was studied cross-sectionally from the viewpoints of somatic growth and reproductive maturation. Distance and velocity curves were expressed using spline function method. Males showed adolescent growth acceleration in body weight, with a peak at 7.86 yrs of age, but not in trunk length. Females showed continuous rapid growth from mid-juvenile to adolescent phase in both body weight and trunk length, but no isolated adolescent spurt. The Sanwa chimpanzees matured at about 12.5 yrs of age for females and 15.0 yrs for males. The mean adult weights and trunk lengths were 53.2 kg and 507.8 mm for males and 42.7 kg and 481.6 mm for females. The Sanwa chimpanzees had similar growth patterns to those of the Yerkes chimpanzees, although they showed a slight delay in infancy, and a higher growth rate from the early juvenile phase onwards. Growth patterns in these two laboratories may be regarded as “normative” for laboratory-reared chimpanzees. They matured earlier than wild chimpanzees by more than two years. The major reason for the retarded maturation in wild chimpanzees is the delay of growth from infant to the early juvenile phases (0–4 yrs of age), probably owing to a limited nutritional supply from the mother. Development of the testes comprised three phases: slow growth from infant to juvenile (until 6.4 yrs); rapid growth around adolescence (until 9.2 yrs); and adult (mean testicular volume, 187 cm³). Setting the nutritional standard at 2,000–2,600 Cal/day (= Kcal/day) per adult, calories were considered for captive chimpanzees in each age class.     

Ontogeny of shell morphology and shell strength of the marine snails Littorina obtusata and Littorina mariae: different defence strategies in a pair of sympatric, sibling species

The sibling marine snails Littorina obtusata (L.) and Littorina mariae Sacchi & Rastelli are sympatrically distributed and the shells of both species are subject to similar breaking forces by predatory crabs. Nevertheless, the two species exhibit rather different growth and defence strategies. To determine growth patterns, we measured changes in five morphological variables with increasing shell length: body whorl thickness at the point of crushing force application, shell height (related to globosity), shell mass, body mass, and apertural lip thickness. We also measured ontogenetic changes in the ability to withstand shell crushing. For most morphological variables, L. mariae showed uniformly allometric growth of juveniles into adults. In contrast, L. obtusata usually exhibited a distinct change in growth pattern upon reaching maturity. As adults, L. mariae showed a more sustained increase in overall shell mass and in body whorl thickness (defence against crushing attacks) and also had proportionally thicker apertural lips (defence against peeling attacks). Littorina obtusata , however, grew to a larger size and their shells could accommodate larger bodies at all sizes. Furthermore, the strength of L. obtusata shells increased faster than could be accounted for by either overall shell mass or thickness at the point of force application, suggesting strengthening by other means such as changes in shell microstructure or shape (other than globosity). These results illustrate the viability of two contrasting antipredator strategies, despite a highly similar phylogenetic history and selective regime.     

Optimal defense strategy: storage vs. new production

If hosts produce defense proteins after they are infected by pathogens, it may take hours to days before defense becomes fully active. By producing defense proteins beforehand, and storing them until infection, the host can cope with pathogens with a short time delay. However, producing and storing defense proteins require energy, and the activated defense proteins often cause harm to the host's body as well as to pathogens. Here, we study the optimal strategy for a host who chooses the amount of stored defense proteins, the activation of the stored proteins upon infection, and the new production of the proteins. The optimal strategy is the one that minimizes the sum of the harm by pathogens and the cost of defense. The host chooses the storage size of defense proteins based on the probability distribution of the magnitude of pathogen infection. When the infection size is predictable, all the stored proteins are to be activated upon infection. The optimal strategy is to have no storage and to rely entirely on new production if the expected infection size n(0) is small, but to have a big storage without new production if n(0) is large. The transition from the "new production" phase to "storage" phase occurs at a smaller n(0) when storage cost is small, activation cost is large, pathogen toxicity is large, pathogen growth is fast, the defense is effective, the delay is long, and the infection is more likely. On the other hand, the storage size to produce for a large n(0) decreases with three cost parameters and the defense effectiveness, increases with the likelihood of infection, the toxicity and the growth rate of pathogens, and it is independent of the time delay. When infection size is much smaller than the expected size, some of the stored proteins may stay unused.     

Plankton predation rates in turbulence: a study of the limitations imposed on a predator with a non-spherical field of sensory perception

This paper presents an extension to previously published work which studied encounter rates of planktonic predators with restricted perception fields, to examine the related problems of prey capture and predation rates. Small-scale turbulence influences planktonic predation in two ways: the extra energy of the flow enhances the number of encounter events between individual predator and prey meso/micro-zooplankton, but it lowers the capture probability (because the time spent by the predator and prey in close proximity is reduced). Typically, an 'encounter' has usually been defined as an event when a potential prey swims (or is advected) to within a distance R of the predator in any direction. However, there is a considerable body of experimental evidence showing that predators perception fields are far from spherical; often they are wedge shaped (e.g. fish larvae), or strongly aligned with the directions of sensory antennae (e.g. copepods); and this is certain to influence optimal predation strategies. This paper presents a theoretical model which for the first time examines the combined problems of both encounter and capture for a predator with a restricted perception field swimming in a turbulent flow. If such a predator adopts a cruising strategy (continuous swimming, possibly with direction changes) the model predictions suggest that predation rates actually vary little with swimming speed, in contrast to predictions made for spherical perception fields. Consequently, cruising predators are predicted to swim at relatively low speeds whilst foraging. However, application of the model to examine the net energy gain of a typical pause-travel predator (the Atlantic cod larva), does predict the existence of an optimal ratio of the length of pauses to time spent swimming (specifically one pause phase to every two travel phases), in line with experimental observations. Kinematic simulations are presented which support these findings.     

Growing through predation windows: effects on body size development in young fish

The degree to which growth in early life stages of animals is regulated via density‐dependent feedbacks through prey resources is much debated. Here we have studied the influence of size‐ and density‐dependent mechanisms as well as size‐selective predation pressure by cannibalistic perch Perca fluviatilis on growth patterns of young‐of‐the‐year (YOY) perch covering several lakes and years. We found no influence of initial size or temperature on early body size development of perch. In contrast, there was a negative relationship between reproductive output and the length of YOY perch at five weeks of age. However, rather than an effect of density‐dependent growth mediated via depressed resources the relationship was driven by positive size‐selective cannibalism removing large individuals. Hence, given a positive correlation between the density of victims and predation pressure by cannibals, size‐dependent interactions between cannibals and their victims may wrongly be interpreted as patterns of density‐dependent growth in the victim cohort. Overall, our results support the view that density‐dependent resource‐limitation in early life stages is rare. Still, patterns of density‐dependent growth may emerge, but from variation in size‐selective predation pressure rather than density as such. This illustrates the importance of taking overall population demography and predatory interactions into account when studying growth patterns among recruiting individuals.