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Materials and methods

LYB (an ear branching selection selected from a common wheat cultivar, Liying No. 3) stably showed abnormal spikes with additional spikelets on extended rachilla. The Chinese Spring monosomic series with normal spikes were used for monosomic analysis of LYB's SS character. Three-five monosomic plants were selected from each the 21 monosomic line, and were artificially pollinated with LYB. At the same time, euploid Chinese Spring was crossed with LYB. A part of F1 monosomic plants and F1 disomic plants was selfed to obtain F2 seeds, other F1 monosomic plants and F1 disomic plants were backcrossed to LYB to obtain corresponding BC1 seeds.

On 3 Nov. 1995, the F1 and F2 seedlings were separately planted. The seeds of BC1 of each combination were sowed in the field. At harvest, all the spikes were observed carefully to determine whether they were SS spikes or normal spikes, and the plants were classified into SS spike type (with at least one SS spike) and normal spike type (with normal spike only). The F2 population which derived from F1 disomic plants was taken as a control in F2 monosomic analysis.


Result and discussion

The F2 monosomic analysis showed that 2A, 2D and 4A F2 populations significantly surpassed the control population in frequency of SS spike plants (
Table 1). The progenies which derived from backcrossing F1 hybrids monosomic for 2A, 2D or 4A to LYB exhibited higher frequency of SS spike plants than other backcross combinations (Table 2). These results mean that chromosomes 2A, 2D and 4A were responsible for SS character.

Koric (1973) found that two genes promoting development of SS and a dominant epistatic inhibitor of SS (Nr) exist in common wheat, and that these genes were independently inherited. The F2 progenies which derived from F1 monosomic plants were homozygous (disomic) or hemizygous (monosomic) corresponding chromosome (s) of LYB. Thus, if a chromosome of LYB carries the gene promoting development of SS, the plants in its corresponding F2 population would be homozygous (disomic) or hemizygous (monosomic) for the gene promoting development of SS. However, according to Koric's conclusion (1973), these plants show SS only when the dominant
Nr gene on other chromosome is absent in their genome. So, the segregation for spike type in the corresponding F2 population would be identical with single gene (Nr) segregation, and the expected segregation ratio would be 1:3 (SS:normal). Only the F2 population which did not possess Nr may show more than 1/4 SS plants. Therefore, to determine whether Nr gene exists or not, chi-square analysis was used to test for the goodness of fit to the segregation ratio of 1:3 in F2 populations. The result showed that F2 progenies of mono-2A and AA fitted the single -gene segregation ratio well (Table 1), and the SS character was recessive. The segregation of monosomics and disomics in 2A and 4A F2 populations could not account for the segregation of spike types, because both disomic and monosomic plants segregated for spike type in these two F2 populations.
As more than two chromosomes were responsible for the SS character in this cross, the segregation ratio of 1:3 (SS:normal) in 2A and 4A F2 populations could not be interpreted as the act of two recessive complementary genes. The most acceptable explanation is that chromosomes 2A and 4A carry genes promoting development of SS, and a dominant strong inhibitor of SS (Nr) exists on other chromosome. Chromosome 2D should be the location of Nr gene, because only it was responsible for the SS character except chromosomes 2A and 4A. The fact that only 2D F2 population showed more than 1/4 SS spike plants supported this conclusion. Obviously, LYB itself could not carry the dominant allele of Nr gene, because it stably showed SS phenotype. Chinese Spring must carry the dominant allele of Nr gene, while LYB should carry its recessive allele(nr) or delete this locus.

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