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Results and discussion

Fertility segregation and utilization of the male sterile gene of Ms2 in triticale
The dominant male sterile Ms2 gene has been located on the short arm of 4D chromosome by Liu and Deng (1982). We have made several crosses and backcrosses between the male sterile triticale Ms2 and hexaploid and octoploid triticale in order to study and utilize its expression in triticale. The values of chi2 in Table 1 indicate that the ratio between the sterile plants and the fertile plants of octoploid (AABBDDRR) Fl is 0.96: 1; the ratio between the sterile plants and the fertile plants of (AABBRRD) Fl of octoploid and hexaploid triticale is 0.91: 1 which all conform to the predicted ratio 1: 1. However, the number of sterile plants significantly decreased according to the increase of backcross number. The ratios were 0.47: 1; 0.45: 1; 0.29: 1 and 0.16: 1 respectively. These ratios are identical with the elimination ratio of 20-50% of D genome chromosome as studied by H. Kihara (Nishiyama 1954). The above results indicate that the dominant male sterile Ms2 gene expressed steadily. The stamens of the sterile plants is abortive, which induces self sterility. The glume opened normally and was able to produce seed after open or artificial pollination, so it is useful to gene recombination. The stamens and pistil of fertile plants develop normally and they are useful to self-fertilize. Therefore, the dominant male sterile triticale is a very useful cross tool for gene recombination and rotational selection.

Transfer of Ms2 male sterile lines
The sterile plants not only play a role in recombination of genes in rotational selection, but also provide half of the genetic factors of hybrids. Nearly all the lines belong to 4-5 grades, because of their narrow genetic base and poor traits, especially poor plumpness. Therefore, we used AH602 AH685, AH999, AH1005 and 20 triticale lines with different traits in transfer breeding before mass crossing to improve the plumpness and other traits of the primary sterile lines. The plumpness had decreased 0.85 class, from 4.52 + or - 0.36 in 1983 to 3.67 + or - 0.54 in 1986 (Table 2), through improvement, and the plumpness of all triticale sterile lines had increased nearly one grade. Some combinations even reached 3.2 grade. Thus they have created good base for rotational selection.

Preliminary results of rotational selection
First, the distribution level of plumpness has increased. It can be seen from the comparison of the mean plumpness of fertile plants in Table 3. C0-C1=0.24, C0-C2=0.72, C-C3=0.80, C1-C2=0.48, C1-C3=0.56, C2- C3=0.08. The number of fertile plants in C1 population averaged 3.43 and it has not exceeded the mean value 3.26 of parents. However, the number of fertile plants in C2 population averaged 2.95 which exceeds the mean value 3.09 of their parents. The plumpness of sterile plants in each rotation (year) has increased by 0.23 which is twice the average conventional lines. In addition, the plumpness proportion of each grade in a population has changed. e.g. the percentage of seeds belonging to 4-5 grades in populations of sterile lines or fertile lines has decreased from 40% to 1%, but the percentage of seeds belonging to grade 3 or above has increased significantly, especially in the population of C2 fertile lines, the percentage of seeds of grade 2 and grade 2.5 has increased from 1% to 15 %. In C3 population the percentage has increased to 20% or more. These results indicate that through rotational selection genes favorable to seed plumpness have increased, thus laying a sound foundation for synthesizing other good traits.

Conclusion
Seed plumpness is a difficult problem in triticale breeding. We have done several thousand of cross combinations and long term pedigree selections, but without good results. Transfer of the dominant male sterile Ms2 gene into octoploid triticale for rotational selection has been made in an attempt to break down the adverse gene linkage with plumpness and to accumulate the favorable quantitative character genes through extensive gene recombination. We attempted to combine good traits such as early maturity, dwarfness, disease resistance and high yield potential on the basis of higher seed plumpness. Rotational selection for seed plumpness improvement is better than pedigree selection and this is related to the basic materials of sterile plants and group combination of recurrent parents. After this work, the mean value of the population was equal to or exceeded that of their parents and showed that it is an effective way for improving seed plumpness of triticale. Further studies are needed in the future because rotational selection has just entered the third rotation.

In the cross and backcross between male sterile octoploid triticale and hexaploid triticale, the proportion of sterile plants was decreased gradually with the increasing number of backcrosses, and this is related to the loss of the D genome chromosome. However, in some combinations there is still a ratio of 1: 1 between sterile plants and fertile plants. It means that 4D chromosome has not been lost or that substitutions and translocations of chromosomes may have occurred. Therefore, the sterile line of hexaploid triticale should be used in rotational selection. It is possible to select new sterile lines of hexaploid triticale from our work, thus a new field will be opened for the cross breeding of hexaploid and octoploid triticales and studies on chromosome engineering of triticale.


References

Liu BH and Deng JY (1982) Genome study and telosomic analysis of the single dominant male-sterile Ta1 gene in common wheat. Scientia Sinica 29: 516-525.

Liu BH and Deng JY (1986) A dominant gene for male sterility in wheat. Plant Breeding 97: 204-209.

Ji FG and Deng JY (1986) Utilization of the dominant male sterile Ta1 gene in triticale breeding programme. 1st Int Triticale Symp, Sydney 77-80.

Darvey NL (1986) Dominant male sterility in hybrid wheat and triticale production. 1st Int Triticale Symp., Sydney 82-85.

Wang CY and Sun YS (1986) Triticale breeding in China, 1st Int Triticale Symp, Sydney 50-58.

Nishiyama I (1954) Cytogenetics in Triticum and its related gene. In: Studies on wheat plants, Ed: Kihara H. Yokendo: 461-471.

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