A fundamental plant evolutionary problem: the origin of land-plant sporophyte; is a new hypothesis possible?
The origin of the sporophyte in land plants represents a fundamental phase in the plant evolution. Today this subject is controversial and, in my opinion, scarcely considered in our textbooks and journals of botany, in spite of its importance. There are two conflicting theories concerning the origin of the alternating generations in land plants: the "antithetic" and the "homologous" theory. These have never been fully resolved. The antithetic theory maintains that the sporophyte and gametophyte generations are fundamentally dissimilar and that the sporophyte originated in an ancestor organism with haplontic cycle by the zygote dividing mitotically rather than meiotically, and with a developmental pattern not copying the developmental events of the gametophyte. The sporophyte generation was an innovation of critical significance for the land-plant evolution. By contrast, the homologous theory simply stated that a mass of cells forming mitotically from the zygote adopted the same developmental plan of the gametophyte, but giving origin to a diploid sporophyte. In this context, a very important question concerns the possible ancestor or ancestors of the land plants. Considerable evidences at morphological, cytological, ultrastructural, biochemical and, especially, molecular level, strongly suggest that the land plants or Embryophyta (both vascular and non-vascular) evolved from green algal ancestor(s), similar to those belonging to the genus Coleochaete, Chara and Nitella, living today. Their organism is haploid for most of their life cycle, and diploid only in the zygote phase (haplontic cycle). On the contrary, the land plants are characterized by a diplo-haplontic life cycle. Several questions are implied in these theories, and numerous problems remain to be solved, such as, for example, the morphological difference between gametophyte and sporophyte (heteromorphism, already present in the first land plants, the bryophytes), and the strong gap existing between these last with a sporophyte dependent on the gametophyte, and the pteridophytes having the gametophyte and sporophyte generations independent. On the ground of all of the evidences on the ancestors of the land plants, the antithetic theory is considered more plausible than the homologous theory. Unfortunately, no phylogenetic relationship exists between some green algae with diplontic life cycle and the land plants. Otherwise, perhaps, it should be possible to hypothesize another scenario in which to place the origin of the alternating generations of the land plants. In this case, could the gametophyte be formed by gametes produced from the sporophyte, through their mitoses or a delayed fertilization process?