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10 Development Interactions Between Clavicipitaleans and Their Host Plants 169<br />

ate impacts of clavicipitalean-produced secondary metabolites on <strong>plant</strong> tissues<br />

themselves. The current scientific wisdom holds that clavicipitalean secondary<br />

metabolites have impacts on animal tissues as feeding deterrents and<br />

other defensive compounds. Whether ergot alkaloids, auxin-like compounds<br />

or other secondary metabolites of these fungi are involved in effecting<br />

changes in <strong>plant</strong> tissues embedded within or adjacent to stromal mycelium<br />

must be further evaluated. It seems likely that this will be a fruitful area for<br />

future investigation.<br />

6.4 Evaporative-Flow Mechanism for Nutrient Acquisition<br />

The stroma of Epichloë maintains a constant flow of water and nutrients into its<br />

mycelium through an evaporation-driven process (White and Camp 1996;<br />

White et al. 1997). Water evaporates rapidly from the <strong>surface</strong> of stromata. As<br />

water evaporates from the stroma it is replaced by water from the <strong>plant</strong>. This<br />

process establishes a flow of water and dissolved nutrients into the stroma from<br />

mycelium interfacing with the vascular bundles and other tissues embedded<br />

within the stroma. Evaporative flow mimics the enhanced transpiration that<br />

occurs in developing inflorescences during elongation of the flowering tillers<br />

of uninfected grasses, but here the stroma is the recipient of the nutrients.<br />

6.5 The Cytokinin Induction Hypothesis<br />

In Atkinsonella hypoxylon stromata form on the inflorescence primordium<br />

and include parts of several leaves as well (Fig. 7). The stromata are gray<br />

(sometimes with areas of a yellow pigment) and produce several different<br />

spore states, including cup-shaped sporodochia that produce moist masses of<br />

ephelidial conidia, and a layer of neotyphodial conidia borne on tips of elongate<br />

conidiogenous cells. It is reasonable to expect that the fungus would<br />

coordinate its development with that of its host grass. For example, the fungus<br />

mycelium must be able to detect when it is growing on an inflorescence primordium<br />

rather than on the tiller meristems. On the tiller meristems it will<br />

produce a low biomass of nonpigmented mycelium and ephelidial conidia,<br />

but no neotyphodial conidia or other structures; while on the inflorescence<br />

primordium the entire suite of morphological structures is produced. One<br />

way for the fungus to coordinate its development to that of the host <strong>plant</strong><br />

would be to use compounds present in the host during different stages of<br />

development as ‘cues’ to initiate developmental stages in the fungus. Our<br />

approach to the search for host compounds that may serve as cues for fungal<br />

development has been a trial and error approach. Over several years, we have<br />

screened hundreds of compounds that might be present in grass tissues to<br />

determine how they affect differentiation of A. hypoxylon.

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