Abstract
The cytological behavior during meiosis of extra constitutive heterochromatin and the effects of such additions to the tomato genome were investigated. The main source of extra heterochromatin was 2S∙2S, a completely heterochromatic isochromosome carrying a nucleolar organizer in each of its arms. Meiotic behavior of a half-heterochromatic, tertiary chromosome 5L∙2S in the presence of 2S∙2S was studied. Wehn 2S∙2S chromosomes were present in more than two doses, achiasmatic simultaneous pairing of three, four and five arms of these chromosomes was observed. "Ectopic" pairing of 2S∙2S with heterochromatin of nonhomologous chromosomes was also observed. It is postulated that "ectopic" pairing neutralized in part the high percentage of anaphase I equational division of the 2S∙2S chromosomes. In some individuals the 2S∙2S chromosome was found organizing an extra, smaller-than-normal nucleolus during meiotic prophase. Conspicuous nucleolar vacuoles were also observed in the progenies of plants carrying 2S∙2S extra chromosomes. It was found that the 2S∙2S chromosome alters diakinesis association and segregation of the 5L∙7S chromosome when both are in the same cell. 2S∙2S chromosomes reduced chiasma frequency, which was also accompanied by reduction of crossing-over for some of the tested gene intervals when 2S∙2S was present in one dose. No effects of 2S∙2S on chromosome pairing of interspecific hybrids were detected. Another effect produced by the addition of 2S∙2S chromosomes was expressed as leaf necrosis in some individuals.