山口 幸

Marine animals show diverse and flexible sexual systems. Here, we review several advancements of theoretical studies made in the last decade. (i) Sex change in coral fishes is often accompanied by a long break in reproductive activity. The delay can be shortened by retaining the inactive gonad for the opposite sex. (ii) Barnacles adopt diverse sexual patterns. The game model was analysed assuming that newly settled larvae choose either growth or immediate reproduction and large individuals adjust male–female investments. (iii) Some parasitic barnacles produce larvae with sexual size dimorphism and others produce larvae with the sex determined after settlement on hosts. (iv) In some fish and many reptiles, sex is determined by the temperature experienced as eggs. The dynamics of sex hormones were studied when the enzymatic reaction rates were followed by the Arrhenius equation. The FMF pattern (male at intermediates temperature; female both at high and low temperatures) required some reactions with enhanced temperature dependence at higher temperatures. The game model provides a useful framework for understanding diverse sexual patterns if we incorporate various constraints, such as unpredictability, cost of trait change and social situations. For further developments, we need to consider constraints imposed by physiological and molecular mechanisms.

In some species of separate sexes, males present a nuptial gift containing nutrition to their mate. Producing a large nuptial gift is a considerable cost to the male, but it may improve his siring success if the female reduces the likelihood to accept another male after receiving a large gift. The female may receive a direct benefit by accepting another male who provides an additional nuptial gift. Additionally, the female may receive an indirect fitness benefit via laying offspring sired by a male who is able to produce a large nuptial gift. We formalized the multivariate quantitative genetics model describing the coevolution of the size of nuptial gift produced by the male (x) and the female's propensity to engage in remating (y). We analyzed the model focusing two cases: [1] remating females receive no indirect fitness benefit, but enjoy direct benefit of nutrition; and [2] remating females receive no direct benefit, but enjoy an indirect fitness benefit due to a positive genetic correlation of x and y, which is possible if random mutations tend to make males produce small nuptial gifts. In both cases, the stable evolutionary equilibrium with neither nuptial gift nor remating (x-=y-=0) always exists. Another stable equilibrium may exist in which male produces nuptial gifts (x->0) and female engage in multiple mating (y->0). We discussed implications to the sexual conflict.

Many marine invertebrates have a benthic adult life with planktonic long feeding larval stages (planktotrophy). In other species, planktonic larvae do not eat, and after a rather short period, they settle and initiate their benthic stages (lecithotrophy). Still other species skip planktonic larval stages altogether, and adults produce benthic offspring (direct development). In this paper, we develop an evolutionary game among different life-cycle types and examine the conditions for each life-cycle type to win in a seasonal environment. The growth rate and mortality of benthic individuals are the same among all three life-cycle types, the local habitat (patches) for benthic individuals have a finite longevity, and adults may engage in a limited dispersal just before breeding. Planktotrophy evolves if the planktonic stages are more efficient in terms of biomass gain than benthic life. Otherwise, lecithotrophy or direct development should evolve. Among them, direct development is more advantageous than lecithotrophy if the cost of having planktonic larvae is large, the habitat for benthic individuals is stable, and adults engage in some dispersal.

Fish live in water with a different osmotic pressure from that in the body. Their gills have chloride cells that transport ions to maintain an appropriate level of osmotic pressure in the body. The direction of ion transport is different between seawater and freshwater. There are two types of chloride cells that specialize in unidirectional transport and generalist cells that can switch their function quickly in response to environmental salinity. In species that experience salinity changes throughout life (euryhaline species), individuals may replace some chloride cells with cells of different types upon a sudden change in environmental salinity. In this paper, we develop a dynamic optimization model for the chloride cell composition of an individual living in an environment with randomly fluctuating salinity. The optimal solution is to minimize the sum of the workload of chloride cells in coping with the difference in osmotic pressure, the maintenance cost, and the temporal cost due to environmental change. The optimal fraction of generalist chloride cells increases with the frequency of salinity changes and the time needed for new cells to be fully functional but decreases with excess maintenance cost.