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The two sexes are defined by the sizes of the gametes they produce, anisogamy being the state with two differing gamete sizes (hence, females and males). The origin of this divergence has received much research interest, both theoretically and empirically. The gamete dynamics (GD) theory is a widely accepted theoretical explanation for anisogamy, and green algae have been an important empirical testing ground for the theory. However, some green and brown algae produce parthenogenetic gametes (gametes that can develop without fusing with another gamete), in contrast to an assumption in GD theory that unfused gametes do not develop. Here, we construct a GD model accounting for parthenogenetic gametes. We find that under conditions of panmixia and highly efficient fertilization (i.e., conditions of classical GD models from 1972 onward), the results remain largely unaltered by parthenogametes. However, under gamete-limited conditions anisogamy evolves less easily in the new model, and a novel result emerges: whereas previous models typically predict the evolution of either anisogamy or small isogamy, the current model shows that large isogamy can evolve when parthenogenetic gametes evolve under conditions of inefficient fertilization. Our analyses uncover unexplored complications relating to sex ratios under this relatively uncharted gametic system. We discuss limitations these complications impose on our models and suggest avenues for future research. We compare model results to algae with parthenogenetic gametes in nature.

There is little information available regarding genomic differences between sexes in ulvophycean green algae. The detection of these differences will enable the development of sex-discriminating molecular markers, which are useful for algae showing little apparent difference between sexes. In this study, we identified male- and female-specific DNA sequences in the ulvophycean marine green alga Monostroma angicava, which has a dioicous heteromorphic haplo-diplontic life cycle, via next-generation sequencing. Fluorescence in situ hybridization (FISH) showed that signals for the sex-specific sequences exist only in the nuclei of the corresponding sex, confirming the specificity of the sequences. Sex-specific molecular markers that targeted these sequences successfully distinguished the sex of gametophytes even in geographically distant populations, indicating that the sex-specific sequences are universal. These results consistently suggest that male and female gametophytes of M. angicava are genetically different, implying that sex may be determined genetically in this alga.

Within-clutch gamete size variability in Monostroma angicava. In many organisms, it is unclear how the size variation in gametes is generated in each clutch (i.e., total gametes produced by a gametophyte for a single spawning) or how gamete size is adjusted. Within-clutch variation in gamete size has been explained as a result of either physiological/developmental constraints or bet hedging during gametogenesis. These two explanations have been assumed to be mutually exclusive, and related observations are conflicting. The slightly anisogamous dioecious green alga Monostroma angicava employs a simple mechanism to produce gametes of each sex: each vegetative cell becomes a single gametangium cell, which synchronously divides to form equally sized gametes. The number of such cell divisions has several variations, which might vary gamete size. We measured the volume of gametangia in each clutch, counted the number of cell divisions in each gametangium and estimated the size of the gametes. We found that larger gametangia divided more times than smaller gametangia in both sexes, although male gametangia were smaller than female gametangia when they underwent the same number of cell divisions. Therefore, the variation in the number of cell divisions during gametogenesis serves to adjust gamete size in each sex rather than to vary it. Within-clutch gamete size variability originates in within-clutch variation in gametangium size: any factors that increase the variation in the size of gametangia can increase the within-clutch variation in gamete size.

The life-cycle system of Ulotrichales, a major order of Ulvophyceae, remains controversial because it is unclear whether the Codiolum phase, a characteristic unicellular diploid generation in ulotrichalean algae, is a zygote or a sporophyte. This controversy inhibits the understanding of the diversified life cycles in Ulvophyceae. To distinguish between zygotes and sporophytes, we have to examine not only whether diploid generations function as sporophytes, but also whether mitosis occurs before meiosis in diploid generations. However, the nuclear behavior in the Codiolum phases is largely unknown, probably because no suitable methods are available. Using fluorescent microscopy with ethidium bromide and transmission electron microscopy of cell-wall-dissected specimens, we report the nuclear behavior in the Codiolum phases of an ulotrichalean alga with a representative life cycle, Monostroma angicava. Each vegetative Codiolum phase had a single polyploid nucleus due to endoreduplication, a type of mitosis without nuclear division. During zoosporogenesis, the nucleus had a structure that would be a meiosis-specific complex. We quantitatively showed that Codiolum phases grew extremely large and produced numerous zoospores. Our results suggest that an event comparable to mitosis occurs before meiosis in the Codiolum phase of M. angicava. This nuclear behavior and the functions (growth and zoospore production abilities) correspond to those of sporophytes. Therefore, the life-cycle system of M. angicava is a heteromorphic haplo-diplontic cycle. This system appears to be widely adopted among other ulotrichalean algae.

In cell divisions, relative size of daughter cells should play fundamental roles in gametogenesis and embryogenesis. Differences in gamete size between the two mating types underlie sexual selection. Size of daughter cells is a key factor to regulate cell divisions during cleavage. In cleavage, the form of cell divisions (equal/unequal in size) determines the developmental fate of each blastomere. However, strict validation of the form of cell divisions is rarely demonstrated. We cannot distinguish between equal and unequal cell divisions by analysing only the mean size of daughter cells, because their means can be the same. In contrast, the dispersion of daughter cell size depends on the forms of cell divisions. Based on this, we show that gametogenesis in the marine green alga, Monostroma angicava, exhibits equal size cell divisions. The variance and the mean of gamete size (volume) of each mating type measured agree closely with the prediction from synchronized equal size cell divisions. Gamete size actually takes only discrete values here. This is a key theoretical assumption made to explain the diversified evolution of isogamy and anisogamy in marine green algae. Our results suggest that germ cells adopt equal size cell divisions during gametogenesis.