28. Distribution of Ef-1 alleles in tropical rice cultivars and common wild rice strains

Kuo-Hai TSAI

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

Gene Ef-1 (abr. E) for early heading was found in various Japanese rice varieties (Tsai 1973, 1986b, 1991; Tsai and Oka 1966; Sato and Hayashi 1985). Recently, this gene was found in certain Indica cultivars (Tsai 1986b) and in a philippine variety of upland Japonica type, Inakupa (Tsai 1986a). Furthermore, similar genes were found in an Indica variety, IR4227 (IR4227-18-3-2) and some Indian Aman varieties (Tsai 1987).

The heading date of IR4227 is about 26 days later than that of T65, but their F1 plants were 5-8 days earlier heading than of T65. The B1F1 of IR4227 with T65 produced segregants like Ef-1 carriers (Tsai 1987). One of such early- heading plants was backcrossed with T65 three times, and an early line (3)IR 4227E was obtained. The F2 of its cross with T65(20)Ea carrying Ef-1a showed heading dates within the parental range (Table 1). There were a few relatively late segregants whose frequency was less than 1/16, but their heading dates were still earlier than that of T65. This suggests that (3)IR4227E and T65(20)Ea have similar alleles at the Ef-1 locus, and their interaction in heterozygotes produces such relatively late-heading segregants.

A long-grain variety from Brazil, BR-IRGA409 shows a heading date about 10 days later than that of T65. A radiation-induced mutant from this variety, N115 showed almost the same heading date as of T65(20)Ea. Their F1 plants were backcrossed with T65 three times, and an early-heading line (3)M15E was fixed. The F2 from T65(20)Ea X (3)M15E showed almost the same heading dates as of the parents (Table 1). This indicates that (3)M15E has an Ef-1 allele.

The Aman varieties from India are highly sensitive to daylength. They produce no panicles in the first crop-season at Taichung and are about 15 days later than T65 in the second crop-season. The reciprocal crosses between T65 and an Aman variety produced F1 plants about 15 days earlier-heading than T65 in the first crop-season and similar to T65 in the second crop-season. In the F2 were found segregants similar to Ef-1 carriers (Tsai 1987). These early-heading plants were backcrossed with T65 five times to obtain (5)Aman-E-3, -4, and -7, which were 2-6 days earlier-heading than T65(20)Ea. When they were crossed with the latter, the F1 showed parent-like phenotypes, and the heading dates of the F2 plants were within the parantal range (Table 2). This indicates that the Aman-E lines have Ef-1 alleles. Genes for photo-sensitivity were also isolated from IR4227-18-3-2 and Aman varieties, and the breeding of isogenic lines with those genes is under way.

An Indica variety from mainland China, Nantsao 33, is about one-month earlier heading than that of T65. This variety was found to carry a late-heading gene ef-4(t) and an early heading gene (Tsai 1991). The early-heading gene was introduced into T65, and three backcrosses of the hybrid produced a semi-isogenic line (3)TK45E. Its cross with T65(20)Ea showed heading-time variations (Table

Table 1. Allelism test for earliness genes found in Indica varieties
==============================================================================
                                      Heading date
Line and F2              ===================================No. of  Mean+/-S.D.
population               Sep.   (1990)                 Oct. plants  (Sep.)
                         7  10  13  16  19  22  25  28  1
==============================================================================
T65 (-T)                                         1   6  1      8    28.O+/-1.60
T65(20)E-                    6   2                             8    10.8+/-1.50
(3)M15E                      2   6                             8    12.5+/-1.24
(3)IR4227E                       9   4                        13    13.9+/-1.35
(3)TK45E                 3   9  12   4   1                    29    12.1+/-2.90
F2:T65(20)Ea x M15E         22  74   4                       100    12.4+/-1.44
F2:T65(20)Ea xIR4227E       12  70  15   2   1               100    13.5+/-1.86
F2:T65(20)Ea xTK45E      6  83  34  11  10  13               157    12.5+/-3.95
==============================================================================
Figures in parentheses show the number of backcrosses repeated.

Table 2.  Allelism test for earliness genes obtained from Aman varieties and
common wild rice (W593, W630 & W1623)
==============================================================================
                                 Heading date                No. of
Line & population     Sep.   (1991)   Oct.                   plants Mean+/-SD
                      16 19 22 25 28  1  4  7 10 13 16 19 22
==============================================================================
T65(-T)                                     4  4  5  5  2  2 22 Oct.13.3+/-4.41
T65(20)Ea                   17  6  2  1                      26 Sept.23.5+/-2.2
(4)W593E15             2  2 10  6  2                         22 Sept.22.4+-3.03
(4)W593E18             5  4  6  2  2  1  1                   21 Sept.21.0+-3.73
F2:T65(20)EaXW593E15   7 20 26 22 16 12  5                  108 Sept.23.9+-4.65
F2:T65(20)EaXW593E18   12 40 63 33 3  5  1                  157 Sept.21.7+-3.13

                         Apr.  (1992)   May
                      23 26 29  2  5  8 11 14 17 20 23 26
T65(-T)                                     2 19  8 10  2    41 May 19.4+/-2.84
T65(02)Ea                       7 16  7  5                   35 May  6.0+/-2.86
(5)AmanE-3                1  7 20  2                         30 May  1.0+/-1.35
(5)AmanE-4                4 15  8  3                         30 Apr.29.0+/-2.11
(5)AmanE-7                1  1 16 15  5  1  1                40 May  4.2+/-3.16
(5)W630E                        6 16  7  1                   30 May  5.1+/-2.27
(4)W1623E                       3 27                         30 May  4.2+/-0.73
F2:T65(20)EaXAmanE-3         4 55 109 17                    185 May  4.1+/-1.90
F2:T65(20)EaXAmanE-4         4 53 113  8 1  1               180 May  4.0+/-1.84
F2:T65(20)EaXAmanE-7      6 98 68 13                        189 Apr.30.4+/-1.58
F2:T65(20)EaXW630E     2 75 49 36 16  2  4                  184 Apr.29.4+/-3.13
F2:T65(20)EaXW1623E       3 59 60 56  3                     181 May  2.2+/-2.32
===============================================================================
1) which suggested the presence of an Ef-1 gene in TK45E.

As mentioned above, rice cultivars so far examined all had Ef-1 alleles. Their wild progenitor, O. rufipogon, was then examined in respect to its genes for heading time. Of three strains used, W593 and W1623 produced no panicles in the first crop-season at Taichung, showing their photoperiod sensitivity. In the second crop-season, they headed several days later than T65. When these wild strains were crossed with T65, the heading dates of the F1 plants were similar to, or a few days earlier than that of T65 (Table 3). In the F2 or B1F1 populations, early-heading plants like Ef-1 carriers were found. These plants were backcrossed with T65 4 to 5 times, and fixed early-heading lines (4)W593E, (5)W630E, and (4)W1623E were obtained. When these lines were crossed with T65(20)Ea, the heading dates of the F2 plants were within the parental range (Table 2). Although some segregants were earlier than the parents, no late-heading segregants like T65 were obtained. The results of these experiments suggest that all the lines have similar alleles at the Ef-1 locus. This indicates the presence of Ef-1 in the wild strains.

Thus, the Ef-1 gene is distributed not only in rice cultivars of Indica and

Table 3.  Mean heading dates of wild-rice strains as compared with that of T65
==============================================================================
Line      Mean heading date   Days to heading of F1    Origin of strain
         ===================  =====================
         1st crop  2nd crop   1st crop  2nd crop
==============================================================================
T65     May 24      Oct. 1          0         0        Taichung 65, Taiwan
W593    No head.        31         +5        +3        Render,Binjai,Malasia
W630    May 27          15          9       -12        Magwe, Burma
W 1623  No head.        21         +4       -10        Pate,Tauyuan,Taiwan
==============================================================================
F1:  Cross with T65.
Japonica types but in their wild progenitors also, so far as the author has examined. Its recessive allele, ef-1, found in T65 is rather a rare variant, which is limited to T65 and other Ponlai varieties of Taiwan that are derivatives of T65. The Ef-1 gene controls the time of floral initiation or the duration of vegetative period, with different effects according to interacting modifiers. Different isoalleles are found at this locus. Probably, as suggested by Tsai (1976), the locus covers a region containing several subloci and within-locus recombination brings about various isoalleles.

References

Sato, Y.I. and K. Hayashi, 1985. The genetic control of basic vegetative phase in early-maturing rice varietice. Jpn. J. Breed. 35: 160-166.

Tsai, K.H., 1973. Comparison of intra-alieleic structure related to the major earliness genes involved in isogenic and radiation-induced early maturing lines of a rice variety, Taichung 65. J. Agric. Assoc. China, N.S. 84: 23-47. (in Chinese with English summary)

Tsai, K.H., 1976. Studies on earliness genes in rice, with special reference to analysis of isoalleles at the E locus. Jpn. J. Genet. 51: 115-128.

Tsai, K.H., 1986a. Genes controlling heading time found in a tropical Japonica variety. RGN 3: 71-73.

Tsai, K.H., 1986b. Genes at the Ef-1 locus found in Fujisaka 5 and two Chinese Indica varieties. RGN 3: 73-74.

Tsai, K.H., 1987. Genes for early heading found in tropical late-heading rice varieties. RGN 4: 85-86.

Tsai, K.H., 1991. Genes for late-heading and their interaction in the background of Taichung 65. Rice Genetics II, p. 211-215. Intern. Rice Res. Inst., P.O. Box 933, Manila, Philippines.