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Introduction to Fungi, Third Edition

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UREDINIOMYCETE YEASTS<br />

667<br />

Fig 24.4 Microscopic features of Phaffia rhodozyma (a) and<br />

Xanthophyllomyces (b,c), a closely related teleomorph.<br />

(a) Vegetatively dividing yeast cells. (b) Thick-walled<br />

chlamydospore in the process of germination bybudding about<br />

3 h after transfer <strong>to</strong> a fresh medium. (c) Aseptate elongate<br />

basidia of Xanthophyllomyces sp. producing basidiospores at<br />

their tips. All images <strong>to</strong> same scale.<br />

complex of several species which are difficult <strong>to</strong><br />

distinguish by any means other than DNA<br />

sequencing (Bai et al., 2002). Rhodo<strong>to</strong>rula spp.<br />

also reproduce vegetatively by budding but are<br />

unable <strong>to</strong> produce ballis<strong>to</strong>conidia. Apart from<br />

this difference, the two genera are very close <strong>to</strong><br />

each other, overlapping in phylogenetic trees<br />

and also possessing similar life cycles.<br />

Life cycle<br />

The budding phase of Sporobolomyces roseus is<br />

uninucleate, and nuclear division occurs at the<br />

time of bud formation (Buller, 1933). If ballis<strong>to</strong>conidia<br />

are <strong>to</strong> be formed, a conical sterigma<br />

develops vertically and bears an asymmetric<br />

spore which strongly resembles a basidiospore<br />

(Figs. 24.1e and 24.5). A daughter nucleus passes<br />

in<strong>to</strong> this spore. Following the fusion of Buller’s<br />

drop with the adaxial blob, the spore is flicked<br />

away for a distance of about 0.1 mm. Colonies of<br />

S. roseus begin <strong>to</strong> form ballis<strong>to</strong>conidia after<br />

2 3 days on most agar media, and they are<br />

recognized by the presence of numerous satellite<br />

colonies outside of the margin of the parent<br />

colony. A single sterigma may produce a second<br />

or even a third spore, and occasionally two or<br />

three sterigmata arise from one cell.<br />

Although S. roseus can form pseudohyphae<br />

and true hyphae, no sexual reproduction has<br />

been observed. However, a related species,<br />

Sporidiobolus (Sporobolomyces, Aessosporon) salmonicolor<br />

(formerly also called S. odorus), does undergo<br />

the full sexual cycle (Fig. 24.7). This species is<br />

heterothallic with a bipolar (unifac<strong>to</strong>rial) mating<br />

system (Fell & Statzell-Tallman, 1981). Following<br />

conjugation between compatible haploid uninucleate<br />

cells, a dikaryotic mycelium with clamp<br />

connections develops and eventually produces<br />

globose binucleate teliospores (Fig. 24.6; Bandoni<br />

et al., 1971). Karyogamy follows. Meiosis occurs<br />

during teliospore germination, which gives<br />

rise <strong>to</strong> an aseptate metabasidium. This buds<br />

off haploid monokaryotic basidiospores at its<br />

tip. Life cycles of this kind have also been<br />

described in some other members of the<br />

Sporidiales, with minor variations such as<br />

the presence of transverse septa in the metabasidia,<br />

or the absence of ballis<strong>to</strong>conidia (see<br />

Fig. 24.1f).<br />

There is, however, a complication in the life<br />

cycle of S. salmonicolor because, according <strong>to</strong><br />

van der Walt (1970), yeast cells may be haploid<br />

or diploid. The diploid cells are larger than the<br />

haploid ones. Both types are capable of reproducing<br />

by budding and by producing ballis<strong>to</strong>conidia,<br />

but only the diploid cells can additionally<br />

develop directly in<strong>to</strong> thick-walled teliospores.<br />

These germinate by means of a short aseptate<br />

promycelium producing yeast-like basidiospores<br />

at its tip (Fig. 24.7).<br />

According <strong>to</strong> Fell and Statzell-Tallman (1998),<br />

a similar life cycle is found in Rhodosporidium<br />

<strong>to</strong>ruloides (anamorph Rhodo<strong>to</strong>rula glutinis). In this<br />

species, diploid yeast cells are thought <strong>to</strong> arise if<br />

there is a failure of meiosis during teliospore<br />

germination. In contrast <strong>to</strong> the observations by

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