SPONTANEOUS IR DUPLICATIONS GENERATED AT MITOSIS IN ASPERGILLUS NIDULANS: FURTHER EVIDENCE OF A PREFERENTIAL SITE OF TRANSPOSED ATTACHMENT

ABSTRACT A radiation-induced translocation, T(IIR → IIIL), has been shown to be nonreciprocal and to have most of IIR, including its terminus, attached uninverted to the terminus of IIIL.—Progeny with the IIR segment in duplicate, obtained from crosses of T(IIR → IIIL) to strains with a standard genome, were unstable at mitosis; like earlier duplication strains, they suffered deletions from either duplicate segment. Frequent mitotic crossing over occurred between the duplicate IIR segments so that, following deletions, more than two classes of stable, balanced products arose from each heterozygous duplication strain.— Spontaneous, mitotically arising duplications of the IR segment, bearing the rate-limiting adE20 allele, can be selected on adenine-free medium on which they emerge as vigorous sectors from the stunted adE20 colony. It was shown previously that most such duplications, when selected from a strain with standard genome, had the terminal IR segment attached to the end of IIR. Selection has now been made from an adE20 strain carrying T(IIR → IIIL), and seven of the 13 independent IR duplications were linked to the III-IIR translocation complex. In three strains analyzed further, the duplicate IR segments, which included the IR terminus, were attached uninverted to the terminus of IIR; the segments of IR were of approximately equal genetic length.—This supports earlier suggestions that there is a preferential site for the initiation of IR duplications and a preferential site, the IIR terminus, for their attachment.

A MEIOTIC UV-SENSITIVE MUTANT THAT CAUSES DELETION OF DUPLICATIONS IN NEUROSPORA

ABSTRACT The meiotic-3 (mei-3) mutant of Neurospora crassa has several effects: (1) When homozygous, it almost completely blocks meiosis and ascospore formation, (2) it is sensitive to UV, (3) its growth is inhibited by histidine and, (4) it increases the instability of nontandem duplications. This was shown for duplications produced by five different rearrangements and was demonstrated by two different criteria. The effects on meiosis and duplication instability are expressed strongly at 25°; the effects on sensitivity to UV and to histidine are expressed strongly at 38.5° but only slightly at 25°. Nevertheless, all four effects were shown to be due to a single gene. mei-3 is not allelic with previously reported UV-sensitive mutants.—Two other results were obtained that are not necessarily due to mei-3: (1) A cross involving mei-3 produced a new unlinked meiotic mutant, mei-4, which is not sensitive to UV or histidine, and (2) a burst of several new mutants occurred in a different mei-3 stock, including a partial revertant of mei-3.—mei-3 has previously been shown to cause frequent complete loss of a terminal duplicate segment, beginning exactly at the original rearrangement breakpoint. Possible mechanisms are discussed by which a UV-sensitive mutant could cause such precise deletions.

Altered instability due to genetic changes in a duplication strain of Aspergillus nidulans

SUMMARYStrains of Aspergillus nidulans with a duplicate segment are mitotically unstable; they produce phenotypically improved variants following deletions in either duplicate segment, and morphologically deteriorated types. The number of variants produced is characteristic of each duplication strain under the same conditions. After ultraviolet treatment two variants, one more stable and the other less stable than the original strain, were selected. Genetic analysis showed that the increased instability in the less stable variant was due to a translocation involving linkage groups V and VIII. The increased stability of the more stable variant was due to a recessive factor (stf–1) located in linkage group VIII. In the homozygous condition this factor also reduces the number of sectors in a diploid strain. The possible genetic mechanisms explaining the instability alterations are discussed.

Mitotic non-conformity inAspergillus: successive and transposable genetic changes

SUMMARYStrains ofAspergillus nidulanswith, a duplicate chromosome segment are mitotically unstable; in addition to phenotypically improved variants, arising following deletions in either duplicate segment, they give morphologically deteriorated types, some with, enhanced stability. In one isolate, deterioration and increased instability were determined by mutation in a duplicate segment; a more stable derivative no longer had this mutation but had one in another linkage group. Another variant, too unstable for analysis, gave derivatives whose single, new mutations were in different linkage groups. It is proposed that deterioration and increased instability result from tandem duplications on either duplicate segment; transposition of these to non-duplicated regions reduces instability. Another 17 variants had a single new mutation each; mutations, possibly clustered, occurred in all linkage groups. In these strains perhaps transposition preceded analysis. Deteriorated variants gave lineages of types with morphological changes caused by further, superimposed mutations. This continued instability is explained as interaction, in fidelity of replication, of non-homologous chromosome segments.Instability inA. nidulansstems from chromosome imbalance. As imbalance is known or suspected in other cases of instability it may be possible to show common mechanisms for apparently diverse phenomena.

Mitotic non-conformity in Aspergillus nidulans: the production of hypodiploid and hypohaploid nuclei

Strains of Aspergillus nidulans with a chromosome segment additional to the normal complement are vegetatively unstable. Previous work suggested that the deletions occurring at mitosis were confined to the unbalanced segments. It has been shown now that deletions, while probably always involving a duplicate segment, may extend beyond it to produce hypohaploids and hypodiploids, respectively, from unbalanced haploid and unbalanced diploid parents.Hypoploids have been proposed tentatively as an explanation for some cases of phenotypic variegation; on this basis it is possible to account for some of the diverse phenomena shown by, for example, position-effect variegation.