A feeling for the superorganism: expression of plant form in the lichen thallus

Abstract

The composite thalli produced by lichen fungi in symbiosis with algae often show structural convergences with plants. Similar overall thallus forms and branching patterns may arise in lichens with very different anatomical construction, indicating the autonomy of the morphological level of organization. Fungal and algal growth and division may be highly integrated within meristem-like morphogenetic zones in many lichens, whereas in others the symbionts may contribute in a less synchronized fashion to the construction of the thallus. Although thallus-level morphology and morphogenesis may be compared with those of plants, ontogeny of the lichen thallus differs fundamentally. Observations of lichen ontogeny illustrate the formation of the thallus by unification of autonomous, primary cellular elements in co-ordinated growth. In land plants and many algae, by contrast, the plant body is the primary structure, the cellular elements of which represent secondary subdivisions. The convergences in form are based on a common mode of nutrition in combination with cell-wall building materials that impart similar structural potential. The photosynthetic apparatus forming the basis of this mode of nutrition is not a convergent feature, however, but a homologous structure that originated in the cyanobacteria and subsequently passed laterally into diverse biological lineages by repeated endosymbioses. With consolidation of these symbioses as eukaryotic algae and plants, the organismal level of organization was repeatedly re-established with increasing degrees of complexity, and morphological convergences were expressed at these new levels. In lichens, by contrast, the symbiosis is not organismally consolidated; morphological expression instead emerges at the superorganismal level.