A Kinesin-14 Motor Activates Neocentromeres to Promote Meiotic Drive in Maize.

Maize abnormal chromosome 10 (Ab10) encodes a classic example of true meiotic drive that converts heterochromatic regions called knobs into motile neocentromeres that are preferentially transmitted to egg cells. Here, we identify a cluster of eight genes on Ab10, called the Kinesin driver (Kindr) complex, that are required for both neocentromere motility and preferential transmission. Two meiotic drive mutants that lack neocentromere activity proved to be kindr epimutants with increased DNA methylation across the entire gene cluster. RNAi of Kindr induced a third epimutant and corresponding loss of meiotic drive. Kinesin gliding assays and immunolocalization revealed that KINDR is a functional minus-end-directed kinesin that localizes specifically to knobs containing 180 bp repeats. Sequence comparisons suggest that Kindr diverged from a Kinesin-14A ancestor ∼12 mya and has driven the accumulation of > 500 Mb of knob repeats and affected the segregation of thousands of genes linked to knobs on all 10 chromosomes.

Fitness Costs and Variation in Transmission Distortion Associated with the Abnormal Chromosome 10 Meiotic Drive System in Maize.

Meiotic drive describes a process whereby selfish genetic elements are transmitted at levels greater than Mendelian expectations. Maize abnormal chromosome 10 (Ab10) encodes a meiotic drive system that exhibits strong preferential segregation through female gametes. We performed transmission assays on nine Ab10 chromosomes from landraces and teosinte lines and found a transmission advantage of 62-79% in heterozygotes. Despite this transmission advantage, Ab10 is present at low frequencies in natural populations, suggesting that it carries large negative fitness consequences. We measured pollen transmission, the percentage of live pollen, seed production, and seed size to estimate several of the possible fitness effects of Ab10. We found no evidence that Ab10 affects pollen transmission, i.e., Ab10 and N10 pollen are transmitted equally from heterozygous fathers. However, at the diploid (sporophyte) level, both heterozygous and homozygous Ab10-I-MMR individuals show decreased pollen viability, decreased seed set, and decreased seed weight. The observed fitness costs can nearly but not entirely account for the observed frequencies of Ab10. Sequence analysis shows a surprising amount of molecular variation among Ab10 haplotypes, suggesting that there may be other phenotypic variables that contribute to the low but stable equilibrium frequencies.

Dynamic Sequence Evolution of a Sex-Associated B Chromosome in Lake Malawi Cichlid Fish.

B chromosomes are extra chromosomes found in many species of plants, animals, and fungi. B chromosomes often manipulate common cellular processes to increase their frequency, sometimes to the detriment of organismal fitness. Here, we characterize B chromosomes in several species of Lake Malawi cichlid fish. Whole genome sequencing of Metriaclima zebra "Boadzulu" individuals revealed blocks of sequence with unusually high sequence coverage, indicative of increased copy number of those sequences. These regions of high sequence coverage were found only in females. SNPs unique to the high copy number sequences permitted the design of specific amplification primers. These primers amplified fragments only in Metriaclima lombardoi individuals that carried a cytologically identified B chromosome (B-carriers), indicating these extra copies are located on the B chromosome. These same primers were used to identify B-carrying individuals in additional species from Lake Malawi. Across 7 species, a total of 43 B-carriers were identified among 323 females. B-carriers were exclusively female; no B chromosomes were observed in the 317 males surveyed from these species. Quantitative analysis of the copy number variation of B-specific sequence blocks suggests that B-carriers possess a single B chromosome, consistent with previous karyotyping of M. lombardoi A single B chromosome in B-carriers is consistent with 2 potential drive mechanisms: one involving nondisjunction and preferential segregation in a mitotic division prior to the germ-line, and the other involving preferential segregation during meiosis I.

Meiotic and mitotic behaviour of B chromosomes of ryegrass / Comportamento dos cromossomos B na mitose e na meiose de azevém

Mitotic and meiotic analyses using conventional and fluorescent stains were employed in plants (accession ETBAZ 055 - origin: Capão do Leão, Rio Grande do Sul State, Brazil) from the Germplasm Active Bank of Ryegrass (Banco Ativo de Germoplasma de Azevém) of Embrapa, for observing the behaviour of B chromosomes. In several meiotic stages, there were observed up to two B chromosomes, which have presented an unstable behaviour regarding their precocious ascension to metaphase I or delays during anaphase I. At the end of the process, the Bs showed predominantly segregation for the nuclei under formation in order to guarantee their propagation. Concerning the cells that comprise the anther tapetum and the root meristems, the B chromosomes have presented a more stable behaviour.

Análises mitóticas e meióticas com coloração convencional e fluorescente foram realizadas em plantas (acesso ETBAZ 055 - procedência Capão do Leão, Rio Grande do Sul) provenientes do Banco Ativo de Germoplasma de Azevém da Embrapa para observar o comportamento dos cromossomos B. Em vários estágios da meiose, foram observados um a dois cromossomos B, os quais apresentaram um comportamento instável, com ascensão precoce na metáfase I ou atrasos na anáfase I. Ao final do processo, os Bs exibiram predominantemente segregação para os núcleos em formação, a fim de garantir a sua propagação. Nas células que constituem o tapete da antera e nos meristemas da raiz, os cromossomos B apresentaram um comportamento mais estável.