第25章種子をつくらない植物

All sexually reproducing organisms have both haploid and diploid cells in their life cycles. In organisms with haplontic life cycles, the haploid stage is dominant, while in organisms with a diplontic life cycle, the diploid stage is the dominant life stage. Dominant in this context means both the stage in which the organism spends most of its time, and the stage in which most mitotic cell reproduction occurs—the multicellular stage. In haplontic life cycles, the only diploid cell is the zygote, which undergoes immediate meiosis to restore the haploid state. In diplontic life cycles, the only haploid cells are the gametes, which combine to restore the diploid state at their earliest convenience. Humans, for example, are diplontic.

Alternation of generations describes a life cycle in which an organism has both haploid and diploid multicellular stages (Figure 25.2). This type of life cycle, which is found in all plants, is described as haplodiplontic.

In alternation of generations, the multicellular haploid form, known as a gametophyte, is followed in the developmental sequence by a multicellular diploid form, the sporophyte. The gametophyte gives rise to the gametes (reproductive cells) by mitosis. This can be the most obvious phase of the life cycle of the plant, as in the mosses, or it can occur in a microscopic structure, such as a pollen grain, in the seed plants. The evolution of the land plants is marked by increasing prominence of the sporophyte generation. The sporophyte stage is barely noticeable in non-vascular plants (the collective term for the plants that include the liverworts and mosses). In the seed plants, the sporophyte phase can be a towering tree, as in sequoias and pines.

Protection of the embryo is a major requirement for land plants. The vulnerable embryo must be sheltered from desiccation and other environmental hazards. In both seedless and seed plants, the female gametophyte provides protection and nutrients to the embryo as it develops into the new sporophyte. This distinguishing feature of land plants gave the group its alternate name of embryophytes.

The sporophyte of seedless plants is diploid and results from syngamy (fusion) of two gametes. The sporophyte bears the sporangia (singular, sporangium). The term “sporangia” literally means “a vessel for spores,” as it is a reproductive sac in which spores are formed (Figure 25.3). Inside the multicellular sporangia, the diploid sporocytes, or mother cells, produce haploid spores by meiosis, during which the 2n chromosome number is reduced to 1n (note that in many plants, chromosome number is complicated by polyploidy: for example, durum wheat is tetraploid, bread wheat is hexaploid, and some ferns are 1000-ploid). The spores are later released by the sporangia and disperse in the environment. When the haploid spore germinates in a hospitable environment, it generates a multicellular gametophyte by mitosis. The gametophyte supports the zygote formed from the fusion of gametes and the resulting young sporophyte (vegetative form). The cycle then begins anew.

Plants that produce only one type of spore are called homosporous and the resultant gametophyte produces both male and female gametes, usually on the same individual. Non-vascular plants are homosporous, and the gametophyte is the dominant generation in the life cycle. Plants that produce two types of spores are called heterosporous. The male spores are called microspores, because of their smaller size, and develop into the male gametophyte; the comparatively larger megaspores develop into the female gametophyte. A few seedless vascular plants and all seed plants are heterosporous, and the sporophyte is the dominant generation.

The spores of seedless plants are surrounded by thick cell walls containing a tough polymer known as sporopollenin. As the name suggests, it is also found in the walls of pollen grains. This complex substance is characterized by long chains of organic molecules related to fatty acids and carotenoids: hence the yellow color of most pollen. Sporopollenin is unusually resistant to chemical and biological degradation. In seed plants, in which pollen is the male gametophyte, the toughness of sporopollenin explains the existence of well-preserved pollen fossils. Sporopollenin was once thought to be an innovation of land plants; however, the charophyte Coleochaetes also forms spores that contain sporopollenin.