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BIOLOGY 21<br />

genus, and the manner of growth can sometimes be altered by Regulating the<br />

environment.<br />

It is now generally accepted that meiosis normally occurs in both families at<br />

the onset of germination, and that segments of the promycelium and the firstformed<br />

sporidia are haploid. The dicaryophytic condition arises by the fusion of<br />

either promycelial cells, sporidia, or hyphae derived from them. Cultural conditions<br />

affect conjugation, and the absence of fusions in any one species may only<br />

indicate that the right conditions have not been found. It is clear, however,<br />

that fusions occur more readily in some species than in others, and that even<br />

physiologic races differ in this res'pect.<br />

DEVELOPMENT OF SPOBIDIA ON THE HOST<br />

In many smut diseases the parasite disappears from view after the initial<br />

infection, and only becomes visible to the eye when sori have developed and<br />

chlamydospores are exposed. In a few species, all members of the Tilletiaceae,<br />

the parasitic mycehum emerges through the stomata or between the epidermal<br />

cells and develops sporidia, sometimes in such profusion that infected organs<br />

are powdery with spores.<br />

The first clear account of this so-called 'conidial stage' was given by Woronin<br />

(1882). Plants of Trientalis europaea infected by Tuburcinia trientalis produce,<br />

after the winter rest, shoots which are white on the lower surface. Woronin<br />

described the sporidiophores, which grow in tufts through the stomata ^nd<br />

between the cells, as non-septate, thin, and bent in such a way that the terminal<br />

sporidia lie horizontally. The sporidia are pyriform, 11-15 (j,, hyahne, with<br />

finely granular protoplasm or a small vacuole. They fall easily and a second<br />

sporidium is produced, but the method of discharge is unknown. If sown on the<br />

surface of a leaf, the germ-tube enters and in 12 to 20 days black flecks, the<br />

young sori, appear. Fusions between sporidia were not observed and the number<br />

of their nuclei is unknown.<br />

Kiihn (1883) germinated the spores of a parasite oi Primula, which he named<br />

Paipalopsis irmischiae, but gave no details as to size or mode of origin of the<br />

spores. Schroeter (1887), listing this as a doubtful member of the Ustilaginales,<br />

stated that the parasite passes through the flowering stem into the floral organs,<br />

forming white powdery spore masses which often fill the whole corolla tube.<br />

The spore, which ha^ a smooth, COIOUEIQSS epispore, is spherical (3-6 /a), and<br />

germinates to form a thin germ-tuKe, the tip of which again forms sporidia. It<br />

is not clear if these Sporidia are like the spores from the corolla tube. Wilson<br />

(1915), recording the fungus from Kent, referred to large numbers of small<br />

unicellular spores present as meal-like masses in the open flower, glueing the<br />

stamens together and partially filling the base of the corolla tube. Viable pollen<br />

was, also present, and Wilson suggested that insects carry spores with poUen to<br />

healthy flowers, but inoculation experiments were unsuccessful. Fusions between<br />

sporidia were observed and, after the passage of one nucleus through the<br />

connecting bridge, the binucleate sporidium developed one or more germ-tubes.<br />

It is thought that members of the genus Thecaphora also form sporidia on stamens<br />

of the host, but no good account of this behaviour has been published (see p. 81,<br />

and Brett, 1940).<br />

Sporidia develop freely on the foliage of plants attacked by some species of

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