All land plants, and some algae, have life cycles in which a haploid gametophyte generation alternates with a diploid sporophyte, the generation of a plant or alga that has a double set of chromosomes. A multicellular sporophyte generation or phase is present in the life cycle of all land plants and in some algae.
The sporophyte produces spores (hence the name), by meiosis. These meiospores develop into a gametophyte. Both the spores and the resulting gametophyte are haploid, meaning they only have one set of homologous chromosomes. The mature gametophyte produces male or female gametes (or both) by mitosis. The fusion of male and female gametes produces a zygote which develops into a new sporophyte. This cycle is known as alternation of generations or alternation of phases.
In the normal course of events, the zygote and sporophyte will have a full double set of chromosomes again. An exception is when a diploid and haploid gamete fuse, resulting in a triploid sporophyte, which will usually be sterile, as dividing three sets of chromosomes into two halves causes complications.
Bryophytes (mosses, liverworts and hornworts) have a dominant gametophyte stage on which the adult sporophyte is parasitic. The embryo of the sporophyte develops from the zygote within the female sex organ or archegonium, and in its early development is therefore nurtured by the gametophyte. Because this embryo-nurturing feature of the life cycle is common to all land plants they are known collectively as the Embryophytes.
Most algae have dominant gametophyte generations, but in some species the gametophytes and sporophytes are morphologically similar (isomorphic). An independent sporophyte is the dominant form in all clubmosses, horsetails, ferns, gymnosperms, and angiosperms (flowering plants) that have survived to the present day. Early land plants had sporophytes that produced identical spores (isosporous or homosporous) but the ancestors of the gymnosperms evolved complex heterosporous life cycles in which the spores producing male and female gametophytes were of different sizes, the female megaspores tending to be larger, and fewer in number, than the male microspores.
During the Devonian period several plant groups independently evolved heterospory and subsequently the habit of endospory, in which single megaspores were retained within the sporangia of the parent sporophyte, instead of being freely liberated into the environment as in ancestral exosporous plants. These endosporic megaspores contained within them a miniature multicellular female gametophyte complete with female sex organs or archegonia containing oocytes which were fertilised by free-swimming sperm produced by windborne miniatuarised male gametophytes in the form of pre-pollen. The resulting zygote developed into the next sporophyte generation while still retained within the pre-ovule, the single large female meiospore or megaspore contained in the modified sporangium or nucellus of the parent sporophyte. The evolution of heterospory and endospory were among the earliest steps in the evolution of seeds of the kind produced by gymnosperms and angiosperms today.
References
- P. Kenrick & P.R. Crane (1997) The origin and early evolution of plants on land. Nature 389, 33-39.
- T.N. Taylor, H. Kerp and H. Hass (2005) Life history biology of early land plants: Deciphering the gametophyte phase. Proceedings of the National Academy of Sciences 102, 5892-5897.
- P.R. Bell & A.R. Helmsley (2000) Green plants. Their Origin and Diversity. Cambridge University Press ISBN 0 521 64673 1