Mechanisms and inheritance of first division restitution in hybrids of wheat, rye, and Aegilops squarrosa

First division restitution (FDR) in intergeneric Triticeae hybrids provides an important meiotic mechanism for the production of amphidiploids without the use of colchicine and similar chemicals. The genetic controls of FDR were investigated by examining microsporogenesis and fertility in F1 hybrids of two- and three-way crosses of durum wheat (Triticum turgidum L. var. durum) cultivars Langdon (LDN) and Golden Ball (GB), 'Gazelle' rye (Secale cereale L.), and one accession (RL5286) of Aegilops squarrosa L. The results from two-way crosses indicated that the first meiotic division varied, depending on the hybrid. GB crossed with Ae. squarrosa developed tripolar spindles and prevented congregation of chromosomes at the equatorial plate. The hybrid of GB with rye had a delayed first division. But, the hybrids of LDN with both Ae. squarrosa and rye had a high frequency of FDR. Analysis from the three-way crosses indicated that inheritance in rye crosses differed from those with Ae. squarrosa. FDR segregated in a 1:1 ratio in the rye cross, suggesting that the FDR is controlled by a single gene from LDN. However, FDR fit a 1:3 ratio in the three-way crosses with Ae. squarrosa. Cytological data suggested that tripolar spindles are a major factor preventing FDR in Ae. squarrosa crosses. Some progenies from the three-way cross with rye had a high frequency of monads that resulted from second division failure of FDR cells.Key words: durum wheat, rye, Aegilops squarrosa, first division restitution, tripolar spindle.

Quantitative genetics of 4x-2x hybrid populations with first-division restitution and second-division restitution 2n gametes produced by diploid parents.

A model is proposed to describe the genetic value of 4x-2x hybrids resulting from crosses between tetraploid genitors and diploid genitors that produce 2n gametes. The model takes into account the genetic consequences of the First Division Restitution (FDR) and Second Division Restitution (SDR) meiosis, particularly on the homozygosity level that 2n gametes contribute to 4x-2x hybrids. As genes can be identical by descent, numerous parameters are needed in the classical approach to describe the inbreeding effects on the mean and variance of 4x-2x hybrids. Using the concept of test value, the model allows a large decrease in the number of required parameters. The model gives the components of genetic variance and usual covariances between relatives using these synthetic parameters. The model is then used to study the efficiency of a recurrent breeding scheme to improve diploid genitors for their combining ability with tetraploid genitors. It appears that, in presence of dominance, ignoring the meiosis pattern will lead to an overestimation of additive variance and then of genetic advance. Some genetic considerations on the differences between FDR and SDR mechanisms lead us to suggest an experimental comparison of their respective advantages and disadvantages for the type of considered recurrent selection. An experimental crossing design is proposed to obtain estimates of the genetic parameters needed for this comparison.

Use of half-tetrad analysis to discriminate between two types of 2n egg formation in a potato haploid

The ratio of the first division restitution (FDR) to second division restitution (SDR) 2n eggs was estimated in 4182t, a haploid (2n = 2x = 24) of Solanum tuberosum L. that produces 2n eggs by the two modes. The segregation of three genes previously mapped relative to their centromeres, Pgm-2 (2.0 cM), Mdh-1 (33.5 cM), and 6-Pgdh-3 (30.1 cM) was analyzed in the tetraploid offspring of a 2x × 4x cross. Based on the segregation of the Pgm-2 locus, 39.7% of the progeny originated from FDR 2n eggs and 60.3% from SDR. Segregation patterns of the two distal loci within the FDR-derived 4x subpopulation indicated that the gene–centromere recombination rate during megasporogenesis was significantly reduced for Mdh-1 when compared with a previous estimate during microsporogenesis. In the SDR-derived 4x subpopulation, the gene–centromere recombination rates for Mdh-1 and 6-Pgdh-3 were not significantly different from previous estimates. Tetraploid progeny generated from one 2x × 4x cross where the 2x parent produces 2n gametes by two modes can be used to make an unbiased comparison of the potential breeding value of FDR and SDR gametes.Key words: potato, megasporogenesis, first division restitution, second division restitution, isozyme.

Occurrence and mechanisms of 2n egg formation in 2x potato

The occurrence of 2n eggs in 381 haploids from six tetraploid parents and in 127 plants representing five diploid wild species was detected using 2x × 4x crosses. Sixty-two percent of the haploids and 24% of the wild-species plants produced 2n eggs. Twenty-six haploids and 17 species plants that gave high seed set in 2x × 4x crosses were examined cytologically to determine the frequency and mechanisms of 2n egg formation. There was significant variation in the frequency of 2n eggs among haploids (7–57%) and among species plants (4.9–57.3%). Five mechanisms of 2n egg formation were identified: synaptic variant (genetically first division restitution); delayed meiotic division (first division restitution and second division restitution); omission of the second division (the prevalent mechanism, second division restitution); irregular anaphase II (second division restitution); and failure of second cytokinesis (second division restitution). 2n eggs can be formed by more than one mechanism within a clone. The occurrence of 2n eggs in wild species and the higher frequency of 2n eggs in haploids than in wild species indicate that sexual polyploidization has been involved in the origin and evolution of polyploid series in potato. The high frequency of 2n eggs in both haploids and diploid wild species allows generation of haploid-species hybrids that produce 2n eggs. These hybrids can then be used in the 2x × 4x and 2x × 2x breeding schemes.Key words: haploids, wild species, 2n gametes, first division restitution, second division restitution.