| MadSci Network: Evolution |
It's a bit difficult to gauge the level of my answer as I don't know how much botany you have had. I'll do my best to make it simple and as brief as possible, but it won't be easy. Of necessity, I will have to invoke some rather esoteric concepts without extensive explanation. The term 'plants' is a bit nebulous. In the broad sense (sensu lato) plants include all of the algae, the fungi, and historically, even the bacteria. In a more strict definition (sensu strictu) plants are limited to the vascular plants (the angiosperms or flowering plants, the gymnosperms or non-flowering plants, and the ferns and their relatives) and the bryophytes (mosses, liverworts, and hornworts). This second, sensu strictu group is sometimes called the "land plants." The land plants and those protists formerly classified as the "green algae" or Chlorophyta are now collective referred to as the Viridaeplantae or Viridiplantae (the green plants). It is the land plants (plants, sensu strictu) that I assume you are referring to. Our knowledge of the early evolutionary history of plants is spotty. The land plants are thought to have evolved from certain members of the protist group formerly called the green algae or chlorophytes. The evidence for this assumption is fairly abundant and consists of biochemical and structural similarities of the chloroplasts; biochemical similarities in the storage and metabolism of carbohydrates; similarities in the formation of the embryo and its protective cells; similarities in the mechanism of cell division; and similar nucleotide (DNA and RNA) sequences, i.e. direct genetic similarities. That the land plants are related to the chlorophytes, especially to a certain group of the former Chlorophyta, the charophytes, is undeniable. Whether the land plants evolved from the charophytes is debatable, but currently not fruitfully so as no viable alternative hypothesis has been put forward. A fairly good "family tree" for land plants may be seen here: http://www.fmnh.helsinki.fi/users/haaramo/Plantae/Viridiplantae.htm This is a fascinating site, by the way, and by all means enjoy it, but... take it with a grain of salt as it seems to be somewhat roguish in terms of its adherence to a strict scientific discipline. The family tree diagram (cladogram) shown there is taken from a reputable biology text. Note that there are two main branches of the land plant tree; the bryophytes (the non-vascular plants). and the vascular plants. Both are believed to have evolved from the charophytes but there is too little fossil evidence to show how they relate to one another, if at all, other than the shared ancestry. Well, now we are getting to the point. Without going into laborious detail about reproductive biology, suffice it to say that these two land plant forms have quite different reproductive life cycles. Both alternate, however, between two generations, the sporophyte (spore-bearing stage) and the gametophyte (gamete-bearing stage). The sporophytes are diploid (have cells with double sets of chromosomes in their nuclei) and the gametophytes are haploid (have cells with single sets of chromosomes in their nuclei). Sporophytes produce spores by meiotic cell divisions (cell divisions that reduce the diploid condition to haploid). Gametophytes produce gametes by mitotic cell divisions (cell divisions that replicate the chromosome complement of the nuclei). The gametes subsequently fuse to reestablish the diploid condition and thus found, through mitotic cell growth and development, the new sporophyte cell line in the form of the early plant embryo. Whew! So much for the similarity between the reproductive life cycles of the vascular plants and the bryophytes. They differ remarkably. In the bryophytes, the gametophyte dominates the life cycle. It is what you see when you look at a moss, for example. The sporophyte generation grows on the gametophyte generation in what could be termed a parasitic relationship if it weren't for the fact that they are the same species. It is a parasite only in the sense that a baby is parasitic on its mother. Sporophytes are visible as the little hairy, non-green things that are sometimes seen projecting out of the moss. See here: http://www.sbs.auckland.ac.nz/info/schools/nzplants/moss_sporophyte.htm In the vascular plants, the sporophyte dominates the life cycle. It is what you see when you look and any plant, such as a tree, a vine, a grass, a flower, or a fern. In most vascular plants, the gametophyte is microscopic and grows (again, like a parasite) on the sporophyte body. It eventually becomes part of the seed in the flowering plants and gymnosperms (the seed plants). Take a look: http://io.uwinnipeg.ca/~simmons/ovule5.htm In the ferns and other fern-like, non-seed plants, the gametophyte is tiny and free living (not dependent on the sporophyte) but is host to the sporophyte embryo that, while it is parasitic on the gametophyte at first, quickly grows to independence and (relatively) large size while the nurturing gametophyte withers beneath it. Here you go:That's what a fern looks like when it's a baby (a sporeling). Here is a fern as a toddler:
Here is another sporeling (I can't get enough of these):
I think that last one is not a fern but a lycopod (club moss), perhaps Selaginella. One thing that seems clear is that the charophyte ancestors of the land plants were dominated by the haploid (i.e., gametophyte) stage of the life cycle. Sporophytes, in the form of multicellular descendants of the zygote, do not exist in the extant representatives of this group. Consequently, both vascular and non-vascular plants have elaborated a sporophyte stage by mitotic intervention at the level of the zygote. So, now you have your answer. That's what is meant by the statement "that in the evolutionary history of plants, gametophyte and sporophyte have switched roles of host and parasite..." Aren't you glad you asked?
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