Takahiro Irie

INTER- AND INTRAPOPULATION VARIATION IN THERMAL REACTION NORMS FOR GROWTH RATE: EVOLUTION OF LATITUDINAL COMPENSATION IN ECTOTHERMS WITH A GENETIC CONSTRAINT

Abstract In ectotherms, lower temperatures in high-latitude environments would theoretically reduce the annual growth rates of individuals. If slower growth and resultant smaller body size reduce fitness, individuals in higher latitudes may evolve compensatory responses. Two alternative models of such latitudinal compensation are possible: Model I: thermal reaction norms for growth rates of high-latitude individuals may be horizontally shifted to a lower range of temperatures, or Model II: reaction norms may be vertically shifted so that high-latitude individuals can grow faster across all temperatures. Model I is expected when annual growth rates in the wild are only a function of environmental temperatures, whereas Model II is expected when individuals in higher latitudes can only grow during a shorter period of a year. A variety of mixed strategies of these two models are also possible, and the magnitude of horizontal versus vertical variation in reaction norms among latitudinal populations will be indicative of the importance of “temperature” versus “seasonality” in the evolution of latitudinal compensation. However, the form of latitudinal compensation may be affected by possible genetic constraints due to the genetic architecture of reaction norms. In this study, we examine the inter- and intrapopulation variations in thermal reaction norms for growth rate of the medaka fish Oryzias latipes. Common-environment experiments revealed that average reaction norms differed primarily in elevation among latitudinal populations in a manner consistent with Model II (adaptation to “seasonality”), suggesting that natural selection in high latitudes prefers individuals that grow faster even within a shorter growing season to individuals that have longer growing seasons by growing at lower temperatures. However, intrapopulation variation in reaction norms was also vertical: some full-sibling families grew faster than others across all temperatures examined. This tendency in intrapopulation genetic variation for thermal reaction norms may have restricted the evolution of latitudinal compensation, irrespective of the underlying selection pressure.

Optimal Growth Pattern of Defensive Organs: The Diversity of Shell Growth among Mollusks

Mollusks show a diversity of shell growth patterns. We develop a model for the dynamic resource allocation to defense organs and analyze it with the Pontryagin maximum principle. A typical optimal growth schedule is composed of the initial phase of soft‐body growth without shell followed by a simultaneous growth of shell and soft body and finally the reproductive phase without growth (simultaneous shell growth). If the defensible predation risk is low or if the cost of defense is high, the optimal strategy is to have no shell (shell‐less growth). If defensible predation pressure or general mortality differs before and after maturation, an additional three strategies, characteristic of the exclusive growth of shell or soft body, can be optimal (sequential shell growth, additional body‐expansion growth, and additional callus‐building growth). These optimal strategies are in accord with the patterns observed for mollusks. In particular, the growth strategies with exclusive growth phase of external shells are preferred when durophagous predation pressure after maturation is higher than that before maturation. This result explains the observation that many tropical gastropods with thickened shell lips spend their vulnerable juvenile phase in sheltered habitats.

Increasing Costs Due to Ocean Acidification Drives Phytoplankton to Be More Heavily Calcified: Optimal Growth Strategy of Coccolithophores

Ocean acidification is potentially one of the greatest threats to marine ecosystems and global carbon cycling. Amongst calcifying organisms, coccolithophores have received special attention because their calcite precipitation plays a significant role in alkalinity flux to the deep ocean (i.e., inorganic carbon pump). Currently, empirical effort is devoted to evaluating the plastic responses to acidification, but evolutionary considerations are missing from this approach. We thus constructed an optimality model to evaluate the evolutionary response of coccolithophorid life history, assuming that their exoskeleton (coccolith) serves to reduce the instantaneous mortality rates. Our model predicted that natural selection favors constructing more heavily calcified exoskeleton in response to increased acidification-driven costs. This counter-intuitive response occurs because the fitness benefit of choosing a better-defended, slower growth strategy in more acidic conditions, outweighs that of accelerating the cell cycle, as this occurs by producing less calcified exoskeleton. Contrary to the widely held belief, the evolutionarily optimized population can precipitate larger amounts of CaCO3 during the bloom in more acidified seawater, depending on parameter values. These findings suggest that ocean acidification may enhance the calcification rates of marine organisms as an adaptive response, possibly accompanied by higher carbon fixation ability. Our theory also provides a compelling explanation for the multispecific fossil time-series record from ∼200 years ago to present, in which mean coccolith size has increased along with rising atmospheric CO2 concentration.

Phenotypic Plasticity and Sexual Dimorphism in Size at Post-Juvenile Metamorphosis: Common-Garden Rearing of an Intertidal Gastropod With Determinate Growth

Proximate factors of the intraspecific variation in molluscan shell morphology have long received attention in biology. The intertidal gastropod Monetaria annulus (Mollusca; Gastropoda; Cypraeidae) is particularly suitable for the study of variation in body size, because this species is a determinate grower in the sense that soft-body size shows no further increase after the juvenile stage. Cross-sectional field surveys on post-juvenile individuals have indicated that the mean body size varies widely among populations and is larger in females than in males within populations. To examine whether these patterns are due to genetic differences, we conducted a common-garden rearing experiment with juvenile individuals collected from two populations on Okinawa Island. After adjusting for among-individual differences in initial degree of development, statistical analyses revealed that this species exhibits female-biased sexual size dimorphism mediated by a longer development time rather than by faster growth rates in females. Although wild individuals show a remarkable size difference between populations, no size difference was found between the populations in the individuals reared in a common-garden condition. This result suggests that the among-population size difference does not have a genetic basis and is caused by phenotypic plasticity based on environmental heterogeneity among habitats.

Thermal dependency of shell growth, microstructure, and stable isotopes in laboratory-reared Scapharca broughtonii (Mollusca: Bivalvia)

We experimentally examined the growth, microstructure, and chemistry of shells of the bloody clam, Scapharca broughtonii (Mollusca: Bivalvia), reared at five temperatures (13, 17, 21, 25, and 29°C) with a constant pCO2 condition (∼450 μatm). In this species, the exterior side of the shell is characterized by a composite prismatic structure; on the interior side, it has a crossed lamellar structure on the interior surface. We previously found a negative correlation between temperature and the relative thickness of the composite prismatic structure in field-collected specimens. In the reared specimens, the relationship curve between temperature and the growth increment of the composite prismatic structure was humped shaped, with a maximum at 17°C, which was compatible with the results obtained in the field-collected specimens. In contrast, the thickness of the crossed lamellar structure was constant over the temperature range tested. These results suggest that the composite prismatic structure principally accounts for the thermal dependency of shell growth, and this inference was supported by the finding that shell growth rates were significantly correlated with the thickness of the composite prismatic structure. We also found a negative relationship between the rearing temperature and δ18O of the shell margin, in close quantitative agreement with previous reports. The findings presented here will contribute to the improved age determination of fossil and recent clams based on seasonal microstructural records.

Water-Quality Variables across Sekisei Reef, A Large Reef Complex in Southwestern Japan.

Abstract: At Sekisei Reef in southwestern Japan (24° N), coral cover dramatically decreased in the mid-1980s, probably due to a population outbreak of the coral predator Acanthaster planci. Coral communities subsequently recovered well outside the semiclosed lagoon, but recovery has been poor inside it. Hence, water-quality degradation including eutrophication has been a concern inside the lagoon. In addition, temporal variation in eutrophication parameters is common among high-latitude coral reefs, resulting in difficulties in evaluating them. Therefore, to address these issues, we monitored temperature, salinity, turbidity, chlorophyll-a, NO x -N (NO3-N + NO2-N), and NH4-N concentrations year-round across the lagoon at Sekisei Reef. Turbidity and NO x -N concentration increased with increasing wind velocity, suggesting that variation in turbidity and NO x -N concentrations was attributed to resuspension of bottom sediments, and NO x -N release through regeneration processes of micro-organisms from the sediments and reef frameworks, respectively. In contrast, variation in chlorophyll- a and NH4-N concentrations appears to be mainly controlled by the seasonality of temperature and irradiance. Long retention time of seawater inside the lagoon seems to have enhanced NH4-N assimilation and increase of phytoplankton during summer. Inside the lagoon, turbidity, NO x -N, and summer chlorophyll-a concentrations were higher, and variation in temperature was larger than outside it. Although water quality appears not to be seriously degraded, multiple effects of these water-quality variables might have negatively affected recovery of coral communities inside the lagoon. Recent expansion of land use on nearby islands might have contributed to water-quality degradation inside the lagoon.