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160<br />

James F. White Jr. et al.<br />

4.3 Parasitism of Grass Meristematic Tissues<br />

On the meristematic tissues of grasses, wounds created by insects are unnecessary<br />

since the tissues of meristems, such as the inflorescence primordia, are<br />

bathed in sucrose. Atkinsonella hypoxylon illustrates this point. Atkinsonella<br />

hypoxylon grows superficially on young leaves of grasses as an epiphyte, perhaps<br />

degrading wax in the cuticle to obtain nutrients for epiphytic growth<br />

(White et al. 1991). When the grass begins to produce an inflorescence primordium,<br />

sucrose is mobilized into the primordium to provide energy for its<br />

development. The primordium is surrounded by nutrients in liquid, which in<br />

turn is surrounded by layers of developing leaves. It is believed that the sudden<br />

increase in sucrose availability and concentration triggers rapid mycelial<br />

growth that eventually results in formation of the stroma (White and Chambless<br />

1991). It is interesting that leaves and inflorescence primordia within the<br />

stroma never develop a cuticular layer that would impede flow of nutrients<br />

and moisture to the fungus. Prevention of cuticular development and prevention<br />

of maturation of the inflorescence primordial tissues may be a growth<br />

regulator effect, although they have not yet been identified for this species.<br />

5 Developmental Differentiation of Endophytic and<br />

Epiphyllous Mycelium<br />

5.1 Plant Cell Wall Alteration<br />

Epichloë sp. illustrate many of the physiological capacities needed by<br />

Clavicipitaceae to colonize grasses (White et al. 1991). Epichloë inhabits leaf<br />

sheaths and growing tillers of grass <strong>plant</strong>s. Endophytic mycelium is largely<br />

nonbranched and exclusively intercellular (Fig. 1) and often seen to adhere<br />

closely to parenchyma cell walls as if attached by glue. This may be due to the<br />

partial degradation of cell wall components by the endophytic mycelium.<br />

While it is possible that cell walls are modified by endophytes, they remain<br />

largely intact as evidenced by electron microscopic studies. It is notable that<br />

during stroma development, profound changes have been observed in cell<br />

walls of the grass epidermis.Walls of epidermal cells appear to lose structural<br />

integrity with mycelium of the endophyte frequently penetrating the wall<br />

(Fig. 2).<br />

5.2 Endophytic Mycelial Growth<br />

Endophytic mycelium in young leaves or elongating tillers is frequently narrow<br />

(1 mm across), straight, oriented parallel to the axis of expansion of the<br />

cells and <strong>plant</strong> organ (Fig. 3). Sometimes in very young tissues, the hyphae

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