Development of thallus axes in Usnea longissima (Parmeliaceae, Ascomycota), a fruticose lichen showing diffuse growth

Abstract

While cell wall thickening in plants is generally associated with tissue maturation, fungal tissues in at least two lichens continue to grow extensively while accumulating massively thickened cell walls. We examined Usnea longissima to determine how diffuse growth shapes morphological and anatomical development of thallus axes and how the highly thickened cell walls of the central cord behave in diffuse growth. Fresh material was examined with light and epifluorescence microscopy and conventional and low-temperature SEM. Fixed material was embedded in Spurr's resin, microtome-sectioned, and examined with TEM and light microscopy. Main axes consisted essentially of bare medullary cord tissue; their characteristic morphology developed by destruction of the overlying cortex and consequent stimulation of lateral branch formation. Fungal cells of the cord tissue continually deposited wall layers of electron-transparent substances and layered, electron-dense materials that include UV-epifluorescent components. Discontinuities were evident in the outermost layers; new branch cells grew through wall materials accumulated by older neighboring cells. Sustained diffuse growth of cord tissue in U. longissima underlies the structural transformation of a corticated thallus branch into a long axis. In the cord tissue, diffuse growth may be responsible for the increasingly disrupted appearance of the older, electron-dense cell wall layers, while new wall materials are laid down adjacent to the protoplast. Cell and tissue development appeared comparable to that observed previously in Ramalina menziesii, although accumulation of wall material was somewhat less extensive and with a greater proportion of electron-dense/UV-epifluorescent components.