Evolution of life cycles

All multicellular plants have a life cycle comprising two generations or phases. One is termed the gametophyte, has a single set of chromosomes (denoted 1N), and produces gametes (sperm and eggs). The other is termed the sporophyte, has paired chromosomes (denoted 2N), and produces spores. The gametophyte and sporophyte may appear identical – homomorphy – or may be very different – heteromorphy. The pattern in plant evolution has been a shift from homomorphy to heteromorphy. The algal ancestors of land plants were almost certainly ‹See Tfd›haplobiontic, being haploid for all their life cycles, with a unicellular zygote providing the 2N stage. All land plants (i.e. embryophytes) are ‹See Tfd›diplobiontic – that is, both the haploid and diploid stages are multicellular.[22] Two trends are apparent: bryophytes (liverworts, mosses and hornworts) have developed the gametophyte, with the sporophyte becoming almost entirely dependent on it; vascular plants have developed the sporophyte, with the gametophyte being particularly reduced in the seed plants. There are two competing theories to explain the appearance of a diplobiontic lifecycle. The interpolation theory (also known as the antithetic or intercalary theory)[23] holds that the sporophyte phase was a fundamentally new invention, caused by the mitotic division of a freshly germinated zygote, continuing until meiosis produces spores. This theory implies that the first sporophytes bore a very different morphology to the gametophyte they depended on.[23] This seems to fit well with what is known of the bryophytes, in which a vegetative thalloid gametophyt is parasitised by simple sporophytes, which often comprise no more than a sporangium on a stalk. Increasing complexity of the ancestrally simple sporophyte, including the eventual acquisition of photosynthetic cells, would free it from its dependence on a gametophyte, as seen in some hornworts (Anthoceros), and eventually result in the sporophyte developing organs and vascular tissue, and becoming the dominant phase, as in the tracheophytes (vascular plants).[22] This theory may be supported by observations that smaller Cooksonia individuals must have been supported by a gametophyte generation. The observed appearance of larger axial sizes, with room for photosynthetic tissue and thus self-sustainability, provides a possible route for the development of a self-sufficient sporophyte phase.[23] The alternative hypothesis is termed the transformation theory (or homologous theory). This posits that the sporophyte appeared suddenly by a delay in the occurrence of meiosis after the zygote germinated. Since the same genetic material would be employed, the haploid and diploid phases would look the same. This explains the behaviour of some algae, which produce alternating phases of identical sporophytes and gametophytes. Subsequent adaption to the desiccating land environment, which makes sexual reproduction difficult, would result in the simplification of the sexually active gametophyte, and elaboration of the sporophyte phase to better disperse the waterproof spores.[22] The tissue of sporophytes and gametophytes preserved in the Rhynie chert is of similar complexity, which is taken to support this hypothesis