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Introduction to Fungi, Third Edition

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324 HYMENOASCOMYCETES: PYRENOMYCETES<br />

Fig12.6 Schematic diagram of ascus development in the two pseudohomothallic ascomycetes Neurosporatetrasperma and Podospora<br />

anserina (based on Raju & Perkins,1994). Neurospora tetrasperma: (a,b) First meiotic nuclear division M I<br />

.The mating type alleles A and<br />

a are closely linked <strong>to</strong> the centromere and segregate at the first division. (c) During the second meiotic nuclear division M II<br />

the<br />

spindles of the dividing nuclei lie in tandem parallel <strong>to</strong> the long axis of the ascus. (d) Interphase of M II<br />

; the nuclei are aligned in pairs.<br />

(e) Telophase of a post-meiotic mi<strong>to</strong>sis PMM I<br />

; the spindles are paired and lie transversely <strong>to</strong> the long axis of the ascus. (f) Interphase<br />

of PMM I<br />

; the four pairs of nuclei are realigned more or less regularly <strong>to</strong> one side of the ascus prior <strong>to</strong> ascospore delimitation.<br />

(g) Young ascospores are binucleate and heterokaryotic, containing both kinds of mating type allele. (h) Following a second<br />

post-meiotic mi<strong>to</strong>sis PMM II<br />

, the ascospores contain four nuclei, two of each mating type. Podospora anserina: the alleles for mating<br />

type are not tightly linked <strong>to</strong> the centromere and do not segregate at M I<br />

(b) but at M II<br />

(c). (d f) as for N. tetrasperma.(g)Following<br />

ascospore delimitation, four binucleate, heterokaryotic ascospores are formed. (h) A second post-meiotic division has occurred.<br />

Three of the four resulting nuclei, two of one mating type and one of the other, remain in the body of the ascospore whilst the<br />

fourth nucleus migrates <strong>to</strong> the basal appendage and degenerates.<br />

division (PMM) brings A and a nuclei close<br />

<strong>to</strong>gether in four pairs, and walls develop<br />

around the nuclei and cy<strong>to</strong>plasm so that the<br />

binucleate ascospores each contain one A and<br />

one a nucleus. In P. anserina, the same result is<br />

achieved by different means. The mating type<br />

idiomorphs (þ) and ( ) are sufficiently distant<br />

from the centromere for a single cross-over <strong>to</strong><br />

occur between the centromere and the locus of<br />

the idiomorphs. Second-division spindles do not<br />

overlap and, by the mechanism outlined above,<br />

nuclei of opposite mating types are brought close<br />

<strong>to</strong>gether at the PMM stage and become enclosed<br />

in walls as binucleate ascospores. A further<br />

mi<strong>to</strong>sis brings the number of nuclei in each<br />

ascospore <strong>to</strong> four, and three nuclei remain in the<br />

body of the ascospore whilst one migrates <strong>to</strong> the<br />

tail (Raju & Perkins, 1994).

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