Ascoycete spore killers: Chromosomal elements that distort genetic ratios among the products of meiosis

Raju, NB,  Mycologia,  86:461-473. 1994.

Spore killers (Sk), studied most extensively in Neurospora, are also known in Podospora, Gibberella and Cochliobolus. Spore killers are no doubt present in natural populations of other fungi. Criteria are outlined here for recognizing their presence and distinguishing them from other causes of ascospore death. Killing occurs when one parent carries the killer element (Sk(K)) and the other carries the sensitive counterpart (Sk(S)). When heterozygous, every ascus contains four normal-sized, viable ascospores and four ascospores that are tiny, undeveloped, and inviable. Spore killers are expressed postmeiotically causing the death of ascospores that do not receive the killer element and resulting in gross distortion of allele ratios for Sk-linked genes. There is little or no ascospore death in homozygous killer x killer or sensitive x sensitive crosses. Sk(K) is centromere-linked in linkage group III of Neurospora crassa and N. intermedia. When Sk is heterozygous, crossing over is blocked in a 30 map unit region that spans the centromere. A sensitive nucleus that would otherwise die is rescued, unchanged, if a killer nucleus is also included in the same ascospore. This has been shown for a developmental giant-ascospore mutant of N, crassa and for the naturally heterokaryotic ascospores of N. tetrasperma and Podospora anserina. As with other segregation-distorters in animals and plants, fungal Spore killers have been found only among strains from nature. Spore killers are fairly common in N. sitophila and Gibberella fujikuroi, but none have been found among natural isolates of N. crassa. In N. intermedia, most strains are sensitive to killing and killers are extremely rare. Some neutral strains are found in nature. These neither kill nor are killed. Resistance to killing is conferred by loci linked to the Sk region. The significance of fungal Spore killers for the individual organism and for populations is discussed.