Thallus Differentiation in the Marchantialean Liverwort Asterella wilmsii (Steph.) with Particular Reference to Longitudinal Arrays of Endoplasmic Microtubules in the Inner Cells

This light and electron microscope study of the liverwort Asterellareveals that, as in other Marchantiales, the cells lining thedorsal air chambers are highly vacuolate with numerous amylochloroplastsin the peripheral cytoplasm. The ventral parenchyma in the midribof the thallus contains aseptate fungal hyphae surrounded byan interfacial matrix and host cytoplasm forming transvacuolarstrands. These are lined with microtubules, rarely seen in otherfungal-hepatic associations or in mycorrhizas. Numerous lipidbodies found in all the thallus cells are thought to be associatedwith perennation during the winter dry season. Elongated, thick walled inner thallus cells, between the dorsalair chambers and the fungus-containing tissue, have a cytologicalorganization not previously recorded in land plants. Initiallyhighly vacuolate, with numerous microtubules of random orientationlining the tonoplast, these cells subsequently show interdigitationof vacuoles and cytoplasm producing a labyrinth of sphericaland elongate tonoplast profiles lined by longitudinal arraysof microtubules. At the same time the cytoplasm becomes increasinglyelectron-lucent and the ribosomes, progressively lost from theER, clump together. At maturity the inner thallus cells arehighly polarized with most of the vacuoles lying nearer thethallus apex. In pits in the end walls, numerous plasmodesmata,with expanded cytoplasmic annuli recall the plamodesmatal fieldsin the mesophyll and phloem of the leaves in vascular plants. Far from being supporting parenchyma or sclerenchyma as assumedhitherto, the inner thallus cells of Asterella are clearly highlydifferentiated. Their vacuole microtubule associations are highlysuggestive of a microtubule-based translocation system akinto that seen in many animal cells and perhaps fungal hyphae,but very different from bulk flow in sieve elements and actin-basedcytoplasmic streaming.Copyright 1994, 1999 Academic Press Cytoskeleton, liverwort, microtubules, plasmodesmata, solute transport