9 Autoregulatory Signals in Mycelial Fungi
The information presented in this chapter reflects on the participation of autoregulatory signals in a wide range of mycelial processes. The various examples of this participation also indicate that the action of autoregulatory signals is combined with ot
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Autoregulatory Signals in Mycelial Fungi
UNAI UGALDE1, ANA BELE´N RODRIGUEZ-URRA2
CONTENTS I. II. III. IV. V. VI. VII.
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Germination . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Colony Morphogenesis . . . . . . . . . . . . . . . . . . . . Asexual Development . . . . . . . . . . . . . . . . . . . . . . Sexual Development . . . . . . . . . . . . . . . . . . . . . . . Dimorphism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
185 186 188 191 195 196 197 198
I. Introduction The mycelium is a colony form which combines the ability to adapt its morphology to the underlying substrate and conditions while maintaining functional unity through interconnectedness. Thus, regions that access nutrientrich substrates may support other’s undertaking exploration or differentiation processes. This display of versatility has led to the realisation that the mycelium is governed by a sophisticated chemo-sensitive system. Many of the elements of this system have probably been already inventoried after many decades of research, but the functional connections between them remain largely unknown. On the one hand, many reported colony functions have been ascribed to as yet unidentified chemical signals. On the other, many thousands of molecules have been purified and characterised, of which only a small proportion have been credited with a biological role. 1 Biochemistry Laboratory II, University of the Basque Country, 1072, 20080 Donostia-San Sebastian, Spain; e-mail: [email protected] 2 Biofungitek, SL, Parque Cientı´fico y Tecnolo´gico de Bizkaia, Ibaizabal Bidea (Edificio 800-2), 48170 Zamudio, Spain
Those signals recognised so far comprise two large categories. Firstly, those directed at relations with other organisms, such as hosts and include elicitors and inhibitors, competitors involving antibiotics and predators where mycotoxins are implicated (for further information see Mycota vols V (2nd edn), X (2nd edn) and XI (2nd edn)). The second category consists of selfdirected signals that formerly fell under the generic term hormones (Gooday 1994). They include pheromones, which facilitate the interaction of compatible gametes, and developmental hormones, which regulate the formation or maintenance of differentiated fungal tissues or proto-organs, such as ascomata or basidiomata. These will be dealt with in Wo¨stemeyer et al. (2016), Dyer et al. (2016), Pelkmans et al. (2016), respectively. In addition to hormones, other intracolonial cues convey information on the status of each cell within the mycelium. They often involve extracellular metabolites and results in adaptations which are advantageous to the colony as a living unit. In the past, these signals have been referred to as autoinducers, autoinhibitors, quorum-sensing factors and morphogens. From a functional viewpoi
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