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Introduction to Fungi, Third Edition

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72 STRAMINIPILA: MINOR FUNGAL PHYLA<br />

within which individual vegetative cells are<br />

contained. For this reason, they are sometimes<br />

referred <strong>to</strong> as ‘slime nets’ (Porter, 1990). The<br />

vegetative cells possess a wall which, uniquely,<br />

is produced from Golgi-derived scales of a<br />

polymer of l-galac<strong>to</strong>se (Dick, 2001a). These<br />

scales are located between the plasma membrane<br />

and the inner membrane of the slime net. The<br />

slime net is delimited by an inner and an outer<br />

membrane and is produced by specialized organelles<br />

termed sagenogens or bothrosomes; the<br />

net membranes are continuous with the plasma<br />

membrane at the sagenogen (Perkins, 1972).<br />

Labyrinthulomycota feed by absorption (osmotrophy)<br />

of nutrients. The nets contain degradative<br />

enzymes which can lyse plant material or<br />

microbial cells. Two orders are distinguished.<br />

4.4.1 Labyrinthulales<br />

Members of this order, especially of the genus<br />

Labyrinthula, can be readily isolated from marine<br />

angiosperms such as Zostera and Spartina or from<br />

seaweed by placing a small piece of one of these<br />

substrata directly on low-nutrient sea water agar<br />

augmented with penicillin and strep<strong>to</strong>mycin<br />

(Porter, 1990). Within a few days, a fine network<br />

of strands can be seen extending over the agar<br />

surface (Fig. 4.6). Labyrinthula spp. can be kept in<br />

monoxenic culture with yeasts or bacteria as<br />

food source. These are presumably lysed by the<br />

enzymes contained in the slime net.<br />

A closer examination shows that the network<br />

consists of branched slime tubes within which<br />

spindle-shaped cells move backwards and<br />

forwards (Fig. 4.7a; see Webster, 2006a).<br />

Movement of a speed up <strong>to</strong> 100 mm min 1 has<br />

been reported and is due <strong>to</strong> a system of<br />

contractile actin-like proteins in the slime net<br />

(Nakatsuji & Bell, 1980). Cells occasionally aggregate<br />

<strong>to</strong> form sporangia containing numerous<br />

round cysts. Following meiosis, eight heterokont<br />

zoospores (Figs. 4.6a, 4.7b) are released by each<br />

cyst. These possess a pigmented eyespot not<br />

found in other types of heterokont zoospore<br />

(Porter, 1990). It is, however, unclear whether<br />

zoospores can establish new colonies (Porter,<br />

1990). Asexual reproduction occurs by division<br />

of spindle cells within the slime net, and<br />

fragments of such a colony can establish new<br />

colonies (Porter, 1972). Further details of the life<br />

cycle appear <strong>to</strong> be unknown at present.<br />

Labyrinthula spp. were implicated as pathogens<br />

in a wasting epidemic of eelgrass (Zostera<br />

marina) at the west coast of North America in the<br />

1930s (Young, 1943; Muehlstein et al., 1991),<br />

causing considerable disturbance <strong>to</strong> the lit<strong>to</strong>ral<br />

ecosystem and collateral damage <strong>to</strong> the<br />

local fisheries industry. However, although<br />

Labyrinthula spp. are still frequently associated<br />

with pieces of moribund Zostera shoots, no<br />

further epidemics seem <strong>to</strong> have occurred since.<br />

Instead, a new species, L. terrestris, has recently<br />

been identified as the cause of a rapid blight of<br />

Fig 4.6 Labyrinthula. (a) Zoospore with<br />

long anterior straminipilous flagellum<br />

and a short posterior whiplash flagellum<br />

with a pointed tip (after Amon &<br />

Perkins,1968). (b d) Portions of colonies<br />

at different magnifications. In (c) spindle<br />

cells are seen in swellings in the slime<br />

tracks.

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