第26章種子植物

Gymnosperms, meaning “naked seeds,” are a diverse group of seed plants. According to the "anthophyte" hypothesis, the angiosperms are a sister group of one group of gymnosperms (the Gnetales), which makes the gymnosperms a paraphyletic group. Paraphyletic groups are those in which not all descendants of a single common ancestor are included in the group. However , the "netifer" hypothesis suggests that the gnetophytes are sister to the conifers, making the gymnosperms monophyletic and sister to the angiosperms. Further molecular and anatomical studies may clarify these relationships. Characteristics of the gymnosperms include naked seeds, separate female and male gametophytes, pollen cones and ovulate cones, pollination by wind and insects, and tracheids (which transport water and solutes in the vascular system).

Gymnosperm seeds are not enclosed in an ovary; rather, they are only partially sheltered by modified leaves called sporophylls. You may recall the term strobilus (plural = strobili) describes a tight arrangement of sporophylls around a central stalk, as seen in pine cones. Some seeds are enveloped by sporophyte tissues upon maturation. The layer of sporophyte tissue that surrounds the megasporangium, and later, the embryo, is called the integument.

Gymnosperms were the dominant phylum in the Mesozoic era. They are adapted to live where fresh water is scarce during part of the year, or in the nitrogen-poor soil of a bog. Therefore, they are still the prominent phylum in the coniferous biome or taiga, where the evergreen conifers have a selective advantage in cold and dry weather. Evergreen conifers continue low levels of photosynthesis during the cold months, and are ready to take advantage of the first sunny days of spring. One disadvantage is that conifers are more susceptible than deciduous trees to leaf infestations because most conifers do not lose their leaves all at once. They cannot, therefore, shed parasites and restart with a fresh supply of leaves in spring.

The life cycle of a gymnosperm involves alternation of generations, with a dominant sporophyte in which reduced male and female gametophytes reside. All gymnosperms are heterosporous. The male and female reproductive organs can form in cones or strobili. Male and female sporangia are produced either on the same plant, described as monoecious (“one home” or bisexual), or on separate plants, referred to as dioecious (“two homes” or unisexual) plants. The life cycle of a conifer will serve as our example of reproduction in gymnosperms.

Pine trees are conifers (coniferous = cone bearing) and carry both male and female sporophylls on the same mature sporophyte. Therefore, they are monoecious plants. Like all gymnosperms, pines are heterosporous and generate two different types of spores (male microspores and female megaspores). Male and female spores develop in different strobili, with small male cones and larger female cones. In the male cones, or staminate cones, the microsporocytes undergo meiosis and the resultant haploid microspores give rise to male gametophytes or “pollen grains” by mitosis. Each pollen grain consists of just a few haploid cells enclosed in a tough wall reinforced with sporopollenin. In the spring, large amounts of yellow pollen are released and carried by the wind. Some gametophytes will land on a female cone. Pollination is defined as the initiation of pollen tube growth. The pollen tube develops slowly, and the generative cell in the pollen grain produces two haploid sperm or generative nuclei by mitosis. At fertilization, one of the haploid sperm nuclei will unite with the haploid nucleus of an egg cell.

Female cones, or ovulate cones, contain two ovules per scale. Each ovule has a narrow passage that opens near the base of the sporophyll. This passage is the micropyle, through which a pollen tube will later grow. One megaspore mother cell, or megasporocyte, undergoes meiosis in each ovule. Three of the four cells break down; only a single surviving cell will develop into a female multicellular gametophyte, which encloses archegonia (an archegonium is a reproductive organ that contains a single large egg). As the female gametophyte begins to develop, a sticky pollination drop traps windblown pollen grains near the opening of the micropyle. A pollen tube is formed and grows toward the developing gametophyte. One of the generative or sperm nuclei from the pollen tube will enter the egg and fuse with the egg nucleus as the egg matures. Upon fertilization, the diploid egg will give rise to the embryo, which is enclosed in a seed coat of tissue from the parent plant. Although several eggs may be formed and even fertilized, there is usually a single surviving embryo in each ovule. Fertilization and seed development is a long process in pine trees: it may take up to two years after pollination. The seed that is formed contains three generations of tissues: the seed coat that originates from the sporophyte tissue, the gametophyte tissue that will provide nutrients, and the embryo itself.

Figure 26.8 illustrates the life cycle of a conifer. The sporophyte (2n) phase is the longest phase in the life of a gymnosperm. The gametophytes (1n)—produced by microspores and megaspores—are reduced in size. It may take more than a year between pollination and fertilization while the pollen tube grows towards the growing female gametophyte (1n), which develops from a single megaspore. The slow growth of the pollen tube allows the female gametophyte time to produce eggs (1n).