V. Molecular Mapping and QTL Analysis
32. 
Mapping QTLs for heading behavior using RI population derived from a cross 
between wild and cultivated rice strains
H.W. Cai1 and H. Morishima2
National Institute of Genetics, Mishima, 411-8544) Japan
Present address: 
1) Japan Forage Seed Association, Nishinasuno Branch, 388-5 Higashiakada, 
Nishinasuno, Tochigi, 329-2742 Japan;
2)15-2 Saiwaicho, Hiratuka, 254-0804 Japan

 
Flowering time is an important trait both for cultivated and wild plants to reproduce themselves in their habitats. Growth duration or days to heading (DTH) in rice can be divided into two phases; basic vegetative phase (BVP) and photoperiod sensitive phase (PSP) (Chang et al. 1969). Several major genes for DTH, BVP and PSP have been identified by many workers.
To identify quantitative trait loci (QTL) for heading date, we used 118 recombination inbred (RI) lines derived from a cross between wild rice 0. rufipogon (W1944) and cultivated rice 0. sativa (Pei-Khu). One hundred forty markers were mapped and employed for QTL analysis. Details of map construction are reported elsewhere (Cai and Morishima submitted). Heading dates were observed in 1995 (F6), 1996 (F7) and 1997 (F7) under natural day length. In addition, in 1996 the same set of materials were also tested in automatic short-day field (11.5 hours). Days to heading observed under short day condition and difference between DTH under short-day and long (natural) day conditions were tentatively regarded as BVP and PSP, respectively. Difference in DTH under natural day-length between 1995 and 1996 was regarded as temperature sensitive phase (TSP), as these two years markedly differed in summer temperature.
DTH, BVP, PSP and TSP were analyzed by simple interval mapping (SIM) and composite interval mapping (CIM) using MQTL software (Tinker and Mather 1995). SIM (0.05 significant level, F = 10.9) revealed three DTH loci and six BVP loci (data not shown). CIM (0.05 significant level, F = 10.9) revealed some more loci for DTH and BVP as shown in Table 1. In addition, four PSP loci were identified. Among DTH loci, qDTH-1 and qDTH-8 are accompanied by BVP and PSP loci closely or within the region flanked by the same markers. These two regions might affect both BVP and PSP genes or carry two linked genes. qDTH-3, qDTH-4-1, qDTH-5-1 and qDTH-l1-1 were accompanied by one BVP gene, respectively. QTL for TSP were not found. Composite interval mapping, by which data taken under multiple environments can be analyzed, enabled us to test interaction between QTL for DTH and year. qDTH-2-2 showed significant interaction with year effect. This locus might be responsible for temperature sensitivity.
Direction of additive gene effect mostly coincided with that predicted by phenotypes of parents. Wild alleles increase DTH, BVP and PSP. Only one BVP locus qBVP-4 showed opposite effect.In addition to the genes for heading behavior detected by conventional genic analysis, marker assisted studies revealed many QTLs (Li et a!. 1995, Xiao et a!. 1996). More recently, Yano eta!. (1997) detected two DTH QTLs with relatively large effect and three with smaller effect based on Nipponbare / Kasalath cross. It seems difficult to compare QTLs we found with the known loci reported by others. qPSP-6 flanked by R2l71 and RZ144 might be Sel, though it should be confirmed through fine linkage mapping.

 
 
 

 
Table 1. QTLs for heading behavior detected using composite interval mapping
 
 
   
Character
QTL
Chr.
Flanking markers
F value
R2 (%)
DTH
qDTH-l
qDTh-2-1
qDTH-2-2
qDTH-3
qDTH-4-1
qDTH-4-2
qDTH-5-l
qDTh-5-2
qDTH-6-l
qDTH-6-2
qDTH-8
qDTH-9-l
qDTh-9-2
qDTH-l1-l
qDTH-1l-2
qDTH-12
1
2
2
3
4
4
5
5
6
6
8
9
9
11
11
12
G393-CD0345
G45-BCD88O
C424-G39
C12-Pgil
G271-C174
R1427-RG214
RG480-C246
BCD1823-Bh2
Est2-Pgi2
Catl-Amp7
G2132-G187
Awn-Estl2
(G385)
RZ14I-Acid2
(CD0365)
G1112-RZ257
20.6
14.5
26.5
55
15.8
21
16.6
28
15.7
13.9
38
31.9
16.5
15.2
19.4
18
5.3
3.8
6.8
13.6
4.1
5.4
4.3
7.2
4.1
3.6
9.6
8.2
4.3
4
5
4.7
BVP
qBVP-l
qBVP-3-1
qBVP-3-2 qBVP-4
qBVP-5-1
qBVP-5-2
qBVP-8
qBVP-11
I
3
3
4
5
5
8
11
RG345-G393
C746-BCD454
G232-Pgil
(G271)
BCD738-Estl3
APAGE1-RG480
G2132-Gl87
RZ14I-Acid2
22.1
14.5
20.2
20
17.3
13.5
11.8
11.4
16.2
11
14.9
14.8
12.9
10.2
9
8.7
PSP
qPSP-1
qPSP-6(Sel?)
qPSP-8
qPSP-11
1
6
8
11
RG345-RAPDB1-l
R2l71-RZ144
G2l32-Gl87
(C6a)
16.3
11
12.6
13.8
12.2
8.4
9.6
10.5
 
References
Cai, H.W. and H. Monshima, 1998. QTL clusters reflect adaptive syndromes of domestication in rice. (submitted to TAG).
Chang, T.T., C.C. Li and B.S. Vergara, 1969. Component analysis of duration from seeding to heading in rice by the basic vegetative phase and the photoperiod-sensitive phase. Euphytica 18: 79-91.
Li, Z., S.R.M. Pinson, J.W. Stansel, W.D. Park, 1995. Identification of quantitative trait loci (QTLs) for head-ing date and plant height in cultivated rice (Oryza sativa L.). Theor. Appl. Genet. 91: 374-381.
Tinker, N.A. and D.E. Mather, 1995. MQTL: software for simplified composite interval mapping of QTL in multiple environments. Journal of Agricultural Genomics 1: Article 2.
Xiao, J., J. Li, L. Yuan and S.D. Tanksley, 1995. Dominance is the major genetic basis of heterosis in rice as revealed by QTL analysis using molecular markers. Genetics 140: 745-754.

Yano, M., Y. Harushima, Y. Nagamura, N. Kurata, Y. Minobe, 1. Sasaki, 1997. Identification of quantitative

trait loci controlling heading date in rice using a high-density linkage map. Theor. App!. Genet. 95:
1025-1032.