Differentiation, growth and morphogenesis: Acetabularia as a model system.

The aim of this paper is to review the present knowledge of the main aspects of differentiation of Acetabularia, a unicellular, eukaryotic organism, and to underline the multiple control pathways modulated by circadian rhythmicity. Growth and morphogenesis are sequentially programmed. Timing of cap differentiation is highly dependent on external conditions. The importance of the sequence of processes is shown by experimental disregulation. The alga is a highly polarized cell, both in morphology and in the relative concentrations of a number of the molecules it contains. Apical cap differentiation is regulated at the post-transcriptional level and could also depend in part on polyamines and on proteolytic activity. Acetabularia displays a number of circadian rhythms (CR). These rhythms form an elaborate biological time structure (also called temporal morphology, or morphology in time as opposed to morphology in space): the distribution in the 24 h cycle of the peaks and troughs of the oscillating functions. The oscillations display fixed relations both with the other functions and with external conditions (such as the transition from dark to light). Interestingly, the CR modulate Acetabularia's development, which is influenced by photoperiod; we present preliminary experiments suggesting that disruption of temporal morphology is deleterious to morphogenesis. Induction of growth and of morphogenesis are totally dependent on blue light. However, blue light receptors in plants arc probably multiple, but we present arguments suggesting that flavin-cytochrome b and the associated KHAM-sensitive molecule are present in Acetabularia plasma membrane and are involved in blue light perception. Agents interfering with different steps of signal perception and transduction show that at least some of these steps are temporally regulated. According to recent experiments from our laboratory, the existence of a redox signalling mechanism appears to be highly probable. The phytohormones (or plant regulators), auxin (indole acetic acid), abscisic acid and ethylene, exert cell-regulatory functions and are involved in Acetabularia differentiation. They also modulate at least some circadian rhythms. Finally, circadian rhythms intervene in differentiation and are proposed to have an integrative function. CONTENTS Summary 1 I. Introduction: the cell cycle and morphology of Acetabularia 2 II. Growth and cap morphogenesis: the developmental programme 3 III. Polarity 5 IV. Temporal morphology 6 V. Induction of growth and cap morphogenesis 9 VI. The plasma membrane 12 VII. Hormones: development and metabolic activity in Acetabularia 12 VIII. Phytohormones receptors and insulin receptors 15 IX. Other possible hormones 16 X. Fundamental role of CR: their intervention in modulating multiple steps in differentiation 16 XI. Conclusions and perspectives 17 Acknowledgements 17 References 17.