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The Questions of Developmental Biology

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quite well with all <strong>of</strong> this "hitchhiking" DNA. Animals also have a significant amount <strong>of</strong> foreign<br />

DNA, but aneuploidy and polyploidy can be developmentally harmful to them. When plants are<br />

aneuploid or polyploid, the consequences can be adaptive. Many flowers found in the florist shop<br />

and the wheat used for bread flour are examples <strong>of</strong> successful polyploids.<br />

Despite these major differences among many plants and animals, developmental genetic<br />

studies are revealing some commonalities between them in the regulation <strong>of</strong> basic molecular<br />

mechanisms <strong>of</strong> patterning, along with evolutionarily distinct solutions to the problem <strong>of</strong> creating<br />

three-dimensional form from a single cell.<br />

*<strong>The</strong> term plant loosely encompasses many organisms, from algae to flowering plants (angiosperms). Recent<br />

phylogenetic studies show a common lineage for all green plants, distinct from the red and brown plants. This new<br />

phylogenetic tree differs from older classification schemes in which the kingdom Plantae consisted <strong>of</strong> multicellular,<br />

photosynthetic plants that develop from embryos protected by tissues <strong>of</strong> the parent plant. While this chapter focuses<br />

primarily on the flowering plants, their developmental strategies are best understood in an evolutionary context.<br />

Plant Life Cycles<br />

<strong>The</strong> plant life cycle alternates between haploid and diploid generations. Embryonic development<br />

is seen only in the diploid generation. <strong>The</strong> embryo, however, is produced by the fusion <strong>of</strong><br />

gametes, which are formed only by the haploid generation. So understanding the relationship<br />

between the two generations is important in the study <strong>of</strong> plant development.<br />

Unlike animals(see Chapter 2), plants have multicellular haploid and multicellular diploid stages<br />

in their life cycle. Gametes develop in the multicellular haploid gametophyte (from the Greek<br />

phyton, "plant"). Fertilization gives rise to a multicellular diploid sporophyte, which produces<br />

haploid spores via meiosis. This type <strong>of</strong> life cycle is called a haplodiplontic life cycle (Figure<br />

20.1). It differs from our own diplontic life cycle, in which only the gametes are in the haploid<br />

state. In haplodiplontic life cycles, gametes are not the direct result <strong>of</strong> a meiotic division. Diploid<br />

sporophyte cells undergo meiosis to produce haploid spores. Each spore goes through mitotic<br />

divisions to yield a multicellular, haploid gametophyte. Mitotic divisions within the gametophyte<br />

are required to produce the gametes. <strong>The</strong> diploid sporophyte results from the fusion <strong>of</strong> two<br />

gametes. Among the Plantae, the gametophytes and sporophytes <strong>of</strong> a species have distinct<br />

morphologies (in some algae they look alike). How a single genome can be used to create two<br />

unique morphologies is an intriguing puzzle.<br />

All plants alternate generations. <strong>The</strong>re is an evolutionary trend from sporophytes that are<br />

nutritionally dependent on autotrophic (self-feeding) gametophytes to the opposite -gametophytes<br />

that are dependent on autotrophic sporophytes. This trend is exemplified by comparing the life<br />

cycles <strong>of</strong> a moss, a fern, and an angiosperm (see Figures 20.2 20.4). (Gymnosperm life cycles<br />

bear many similarities to those <strong>of</strong> angiosperms; the distinctions will be explored in the context <strong>of</strong><br />

angiosperm development.)<br />

<strong>The</strong> "leafy" moss you walk on in the woods is the gametophyte generation <strong>of</strong> that plant (Figure<br />

20.2). Mosses are heterosporous, which means they make two distinct types <strong>of</strong> spores; these<br />

develop into male and female gametophytes. Male gametophytes develop reproductive structures<br />

called antheridia (singular, antheridium) that produce sperm by mitosis. Female gametophytes<br />

develop archegonia (singular, archegonium) that produce eggs by mitosis. Sperm travel to a<br />

neighboring plant via a water droplet, are chemically attracted to the entrance <strong>of</strong> the archegonium,<br />

and fertilization results.* <strong>The</strong> embryonic sporophyte develops within the archegonium, and the<br />

mature sporophyte stays attached to the gametophyte. <strong>The</strong> sporophyte is not photosynthetic. Thus

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