8. Detection of a new semi-dwarfing gene, sd-8(t)

Kuo-Hai TSAI

Department of Agronomy, National Chung-Hsing University Taichung, Taiwan 40227, ROC

During recurrent backcrossing of Taichung 65 (abbr. T65) to early-heading donor parents, Tatung-tsailai (A) and Bozu 5 (B), a line similar to T65, (7)ea was obtained from cross A, and another similar line, (15)eb was obtained from cross B. These T65-like lines both showed a heading time about three days earlier than that of T65 and bred true. Their plant height was similar to that of T65. When they were intercrossed, 7 early-heading offtypes occurred out of 2455 F`2` plants (Tsai 1990). One of the early-heading offtypes (AB-60) produced in its progeny several lines differing in heading time, ranging from "E-type" three to six days earlier-heading than the isogenic line of T65 with gene Ef-1a, and "e'-type" four to six days earlier than T65 carrying ef-1, to a late-heading type 15-25 days later than T65. An e'-type plant, AB-60e'-3 (abbr. e'-3) had a heading time 4 to 9 days earlier than T65 and its plant height was 15-19 cm shorter than that of T65 (Tsai 1990). The F`1` plants between T65 and e'-3 showed intermediate heading time between the parents, indicating partial dominance of the earliness gene of line e'-3. But the height of the F`1` plants was similar to that of T65. The F2 population segregated into 1 e'-3(early): 2 intermediate: 1 T65-like (late) for heading time, and 3 tall (T65-like): 1 short (e'-3 like) for plant height (Table 1).

Table 1.  F`1` and F`2` data for plant height in crosses of T65 with three
          semi-dwarf lines (difference from T65 in parentheses)
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                            F`2` Plant number
            F`1` Mean
                       T65-like  sd-7 like  sd-1 like  Total  Chi2     Expected
Cross      (2nd crop)  97-105 cm 87-97 cm   71-82 cm                    ratio
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               cm
T65xsd-1    88 (-11)     129       282        148       559    1.3      1:2:1
T65xsd-7   100           109        40                  149    0.3        3:1
T65xe'-3    99 (-1)      113        35                  148    0.2        3:1
   (sd-8)
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An early-heading line, T65(20)Ef-1a headed about 6 days earlier than line e'-3, and showed semi-dwarfism similarly as e'-3. The F`1` plants between T65(20)Ef-1a and e'-3 headed three days earlier than T65(20)Ef-1a and were similar to the parents in semi-dwarfism. Their F`2` population segregated into 3 early (Ef-1 type): 1 late (e'-3 type) for heading time, and the plants of e'-3 type segregated further into 3 tall (T65-like): 1 semi-dwarf (e'-3 like). The Ef-1a gene has a pleiotropic effect to reduce plant height (Tsai and Oka 1965). When a plant had ef-1 in place of Ef-1, its height is recovered to the stature of T65. The early-heading segregants of T65(20)Ef-1a x e'-3 were all semi-dwarfs and a 1/4 part of late-heading plants (e'-3 type) were also semi-dwarfs. The occurrence of tall and semi-dwarf plants in the late-heading class suggests that the parental lines have different genes for plant height. The semi-dwarfing gene carried by line e'-3 is denominated as sd-x.

Taichung Native 1 (TN1) is known to have the Dee-geo-woo-gen gene, sd-1 (Suh and Heu 1978; Kikuchi and Ikehashi 1984; linkage group 1). The gene was introduced into T65 by 8 backcrosses and a near-isogenic line T65sd-18 was produced. This line was about 26 cm shorter in height than T65. Its F`1` plants with T65 were 11 cm shorter than T65, and the F`2` segregated into 1 tall: 2 intermediate: 1 short types. The allele of T65, sd-1+ is incompletely dominant over sd-1.

On the other hand, a semi-dwarf mutant line, D65-312 was obtained after X-ray irradiation (40 Kr) of T65 seed. This line was about 16 cm shorter and 3 days earlier-heading than T65 (Tsai 1989). Its F`1` plants with T65 showed a similar height to T65, and the F`2` segregated into 109 T65-like: 40 dwarf plants, giving a good fit to the 3: 1 ratio. The semi-dwarfing gene of this mutant was symboled sd-7. It was located on chromosome 5 (Tsai 1991). Crossing experiments showed that sd-1 and sd-7 were independent, and that homozygotes for both had a much reduced plant height. Mean heading time and plant height of lines with these semi-dwarfing genes are summarized in Table 2.

Table 2.  Mean plant height and days to heading of T65 and its near-isogenic
          lines with a semidwarfing gene (difference from T65 in parentheses)
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Line            Plant height (cm)          Days to heading
             1st crop   2nd crop         1st crop     2nd crop
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T65          102(=O)    99(=O)           127(=O)      88(=O)
T65sd-1`8`    76(-26)   73(-26)          128(+l)      87(-1)
T65sd-7`5`    90(-12)   85(-14)          124(-3)      82(-6)
e'-3(sd-8)    95(-7)    82(-17)          115(-12)     77(-11)
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1st crop: planted in winter, 2nd crop: planted in summer.
To test the allelism of semi-dwarfing genes, crosses were made between e'-3 and T65sd-1`8`, and between e'-3 and T65sd-73. The F`1` plants between e'-3 and T65sd-1 had an intermediate height between T65 and e'-3, and the F`2` plants varied from tall (T65-like) to very short (double dwarf). When plants with intermediate heights were pooled, the F`2` segregation pattern fitted 3 tall: 12 intermediate: 1 short ratio (Table 3). Letting the dwarfing gene of e'-3 be sd-x, the F2 data show that sd-x and sd-1 are independent. Although both genes are recessive, their heterozygotes like the F`1` plants show semi-dwarfism possibly due to some interaction of two recessive genes.

Table 3.  Expected and observed frequencies of F`2` plants in crosses of line
e'-3 with sd-8 and two other semi-dwarf lines

                    a)   e'-3xT65sd-1`8`
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       Genotype            Frequency              No. of plants
  sd-8        sd-7         expected             Expected  Observed
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  +  +        +   +         1/16_ 
                                 | 3/16
  +  sd-8     +   +         2/16_|                 26.4      34
          
  +  +        +   sd-7      2/16_
  +  sd-8     +   sd-7      4/16 |
  +  sd-8   sd-7  sd-7      2/16 | 12/16          105.7      99
sd-8 sd-8     +   sd-7      2/16 |
sd-8 sd-8     +   +         1/16 |
  +  +      sd-7  sd-7      1/16_|
sd-8 sd-8   sd-7  sd-7      1/16  1/16              8.8       8
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Differences between expected and observed numbers did not reach the 5% level
of significance.

                    b)   e'-3 XT65sd-7`3`
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       Genotype            Frequency             No. of plants
  sd-8       sd-7          expected             Expected  Observed
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  +  +       +   +          1/16_
                                 | 5/16
  +  sd-8    +   +          2/16_|                44.7       47

  +  +       +   sd-7       2/16_
  +  sd-8    +   sd-7       4/16 |
  +  sd-8  sd-7  sd-7       2/16 |
                                 | 10/16          89.4       86
sd-8 sd-8    +   sd-7       2/16 |
sd-8 sd-8    +   +          1/16 |
  +  +     sd-7  sd-7       1/16_|
sd-8 sd-8  sd-7  sd-7       1/16    1/16           8.9       10
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Differences between expected and observed numbers did not reach the 5% level
of significance.
Further, line e'-3 with sd-x was crossed with T65sd-7. The F`1` plants were somewhat shorter (by 4 cm) than T65, and the F`2` segregated into 5 tall: 10 intermediate: 1 short classes when plants with intermediate heights were pooled, as shown in Table 3. These experimental results indicate that gene sd-x is independent of both sd-1 and sd-7. It is symboled sd-8 tentatively.

The agronomic characters of these lines are summarized in Table 4. Gene sd-1 shortens the upper internodes rather strongly, but sd-7 and sd-8(t) both shortens the second to fifth internodes moderately. Gene sd-8(t) seems to increase spikelets per panicle.

Table 4.  Character values of T65 and its semi-dwarf lines
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                 1st  crop, 1992               2nd crop, 1991
Character   _______________________________  _________________________________ 
            T65  T65sd-1`9` T65sd-7`4` e`-3  T65  T65sd-1`9` T65sd-7`4`  E`-3
                                      (sd-8)                            (sd-8)
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Length, cm
Plant height 97.O  71.4       81.8     81.1  99.4  68.0        85.1      84.2
Panicle      22.2  20.4       21.5     20.8  22.5  20.0        20.7      19.1
Internode 1  41.8  32.5       41.1     40.1  39.8  25.8        38.1      38.3
    "     2  21.4  12.4       15.1     14.0  21.3  13.8        17.1      16.2
    "     3   9.3   4.4        2.4      4.0  10.0   5.3         6.4       6.5
    "     4   2.7   1.2        0.7      1.0   4.0   1.8         1.0       2.5
    "     5   0.8   0.5        0.4      0.5   1.0   0.7         0.6       0.8
Panicles/plant9.0   7.3        6.7      6.2   8.7   9.9        11.6       8.7
Spikelets/panicle139 119     135      185   144   106         139       162
Spikelet length6.8mm 6.8       6.7      6.4   7.0   6.9         6.8       6.4
   "  width   3.3   3.4        3.5      3.3   3.5   3.5         3.3       3.3
1000 grain wt.25.09 24.9      24.7     18.6  25.4  21.0        22.5      20.6
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References

Kikuchi, F. and H. Ikehashi, 1984. Semidwarfing genes of high-yielding rice varieties in Japan. RGN 1: 93-94.

Suh, H. S. and M. H. Heu, 1978. The segregation mode of plant height in the cross of rice varieties, XI. Linkage analysis of the semi-dwarfness of rice variety "Tongil". Korean J. Breed. 10: 1-6.

Tsai, K. H., 1989. An induced dwarfing gene, sd-7(t), obtained in Taichung 65. RGN 6: 99-101.

Tsai, K. H., 1990. Unusual mode of inheritance of heading time and occurrence of weak plants observed in rice. Japan. J. Breed. 40: 133-146.

Tsai, K. H., 1991. Tight linkage of gene sd-7(t) and d`1` found in a cross of Taichung 65 isogenic lines. RGN 8:104.

Tsai, K. H. and H. I. Oka, 1965. Genetic studics of yielding capacity and adaptability in crop plants. 1. Characters of isogenic lines in rice. Bot. Bull. Acad. Sinica 6: 19-31.