Conducting tissues and phyletic relationships of bryophytes

Internal specialized conducting tissues, if present, are restricted to the gametophytic generation in liverworts while they may occur in both generations in mosses. Conducting tissues are unknown in the anthocerotes. Water-conducting cells (WCCs) with walls perforated by plasmodesma-derived pores occur in the Calobryales and Pallaviciniaceae (Metzgeriales) among liverworts and in Takakia among mosses. Imperforate WCCs (hydroids) are present in bryoid mosses. A polarized cytoplasmic organization and a distinctive axial system of microtubules is present in the highly specialized food-conducting cells of polytrichaceous mosses (leptoids) and in less specialized parenchyma cells of the leafy stem and seta in other mosses including Sphagnum. A similar organization, suggested to reflect specialization in long-distance symplasmic transport of nutrients, also occurs in other parts of the plant in mosses, including rhizoids and caulonemata, and may be observed in thallus parenchyma cells of liverworts. Perforate WCCs in the Calobryales, Metzgeriales and Takakia, and hydroids in bryoid mosses, probably evoked independently. Because of fundamental differences in developmental design, homology of any of these cells with tracheids is highly unlikely. Likewise, putative food-conducting of bryophytes present highly distinctive characteristics and cannot be considered homologous with the sieve cells of tracheophytes.