14. Two genes affecting photoperiod sensitivity in rice

Monton POONYARIT, D. J. MACKILL AND B. S. VERGARA

International Rice Research Institute, P. 0. Box 933, Manila, Philippines

The inheritance of photoperiod sensitivity and critical daylength was studied in four Indica rices (Table 1). The parents, F1 and F2 of the six crosses were planted under natural daylength on April 6-7, 1986, at Los Banos, Philippines (14 deg.N latitude).

A bimodal distribution in F2 progenies was observed in all crosses except PR2/SN, using different cut-off points depending upon the distribution (Table 2). F2 progeny of IR26760-27/NS19 showed a good fit to a 9: 7 ratio (late: early) (Table 2). The F2 of the crosses of PR2 or SN with NS 19 or IR26760-27 showed a good fit to the ratio 3:1 (late: early) (Table 2). The F2 progeny of crosses with NS19 were later flowering compared to the progeny of crosses with IR26760-27. It seems that all the F2 Plants of crosses with NS 19 were photoperiod-sensitive and the differences among these genotypes were due to critical daylength.

Two genes that confer photoperiod sensitivity, one dominant (Se-1) and another recessive (se-2), have been identified in rice (Yu and Yao 1968; Yokoo and Fujimaki 1971; Yokoo et al. 1980). Since the photoperiod sensitivity gene Se-1 is closely linked with the isozyme gene Pgi-2 (Kinoshita 1986), the Pgi-2 alleles were used to determine if any of the genes conferring photoperiod sensitivity were allelic to Se-1. Results indicated that PR2, SN and NS19 all had a dominant gene conferring sensitivity at the Se-1 locus. IR26760-27 has a recessive allele at this locus, but has a dominant gene at another locus independent of Se-1 which also delays flowering. Since NS19 has the recessive allele at the same locus, here designated as se-3(t), it flowers much earlier than PR2 or SN. It appears that Se-3(t), when present with the dominant allele at Se-1, confers strong photoperiod sensitivity and short critical daylength. When tillers of F2 plants of PR2/NS19

Table 1.Parents used in genetic study of photoperiod-sensitivity
________________________________________________________________
                          PSP2   Regression       Proposed 
Parent Origin Duration1   (days) Coefficient3     genotype
________________________________________________________________ 
IR26760
-27     IRRI     145        75    -0.12** se-1se-1Se-3(t)Se-3(t) 
Nam Sa- 
gui 19  Thailand 122       114    -0.26** Se-1Se-1se-3(t)se-3(t) 
Puang 
Rai 2   Thailand 193      >135    -0.64** Se-1Se-1Se-3(t)Se-3(t) 
Sac 
Nau     Vietnam  190      >141    -0.57** Se-1Se-1Se-3(t)Se-3(t)
________________________________________________________________ 
1Seeded 18 June 1986.

2 Photoperiod sensitive phase measured as the difference in days to flower under 14 hour photoperiod and the days to flower under 10 hour photoperiod.

3 Regression of flowering duration on seeding dates for 11 dates of plating (May 14-Oct.17) IRRI.

** Significant at the 1% level of probability.

Table 2. Distribution of days to flowering (seeded 6 April) for parents, F1 and F2 and X2 analysis for goodness of fit to gene models

______________________________________________________________
Cross                         Days to flowering
                     _________________________________________
                     80  90  100 110 120 130 140 150 160 170 
______________________________________________________________ 
IR26760-27/NS19  P1               13   1    
                 P2                            5 10 
                 F1                                       11
                 F2      24   31  42  29  31  15  9   12  58
PR2/IR26760-27   P1    
                 P2               11   3
                 F1
                 F2                2  14  23  19  19  18   7
SN/IR26760-27    P1
                 P2                1  11
                 F1        
                 F2   1                6  22  36  19  11   2
PR2/NS19         P1
                 P2                            1  14
                 F1
                 F2                            8  19  29  23
SN/NS19          P1
                 P2                            5   6
                 F1 
                 F2                        1   3  26  39  22
PR2/SN           P1
                 P2
                 F1
                 F2
______________________________________________________________   
 


______________________________________________________________
Cross                         Days to flowering
                     _________________________________________
                       180  190  200  210  220  230  240  250 
______________________________________________________________ 
IR26760-27/NS19  P1             
                 P2                             
                 F1      1                                 
                 F2     55   36   26    2
PR2/IR26760-27   P1                          1    6
                 P2                    
                 F1                    10    1
                 F2      0   11   34  106   84   36   1
SN/IR26760-27    P1                     2    0    9
                 P2               
                 F1           1   10
                 F2      7   33  105   74   29    6
PR2/NS19         P1                     1
                 P2                            
                 F1           6    6
                 F2     38   74  102   49   29                    
SN/NS19          P1                          1    5
                 P2                           
                 F1           9
                 F2     28   89   92   39   15    4                  
PR2/SN           P1                               8
                 P2                          8    4
                 F1                     1    6    4
                 F2                         64  185   17   3 
______________________________________________________________   


______________________________________________________________
Cross                   F/2          F/2         X2              
                       Range         Ratio 
______________________________________________________________ 
IR26760-27/NS19  P1              
                 P2    86-150         7          1.02                          
                 F1   151-210         9         P>0.30
                 F2   
PR2/IR26760-27   P1   
                 P2   112-179         1          2.55                 
                 F1   171-243         3         P>0.10
                 F2   
SN/IR26760-27    P1  
                 P2    79-170         1          1.29            
                 F1   171-227         3         P>0.20
                 F2  
PR2/NS19         P1  
                 P2   138-174         1          2.52                          
                 F1   175-225         3         P>0.10
                 F2  
SN/NS19          P1  
                 P2   129-174         1          0.02                        
                 F1   175-228         3         P>0.90
                 F2  
PR2/SN           P1  
                 P2  
                 F1  
                 F2  
______________________________________________________________   

Fig. 1. Distribution of critical daylength in F2 populations of two crosses.

and SN/NS19 divided during the basic vegetative phase were treated under 5 photoperiods, a bimodal distribution was observed (Fig. 1). The F2 distribution showed a good fit to a 3: 1 ratio indicating that this dominant gene conferred short critical daylength.

Yokoo and Kikuchi (1977) found evidence that different alleles of the Se-1 (Lm) locus control flowering date. A study by Li (1970) suggested that a single gene controlled critical daylength. In the recent study, a single gene independent of Se-1, designated as Se-3(t), affected critical daylength. lt is not clear if this gene is the same as E/1, which also affects photoperiod sensitivity and is independent of Se-1 (Yamagata et al. 1986).

Acknowledgement. The authors express their gratitude to Drs. E. Gtiiderdoni and J. C. Glaszmann, and Mr. B. de los Reyes for helpful comments and assistance in the isozyme studies.

References

Kinoshita, T., 1986. Standardization of gene symbols and linkage maps in rice. In Rice Genetics, p. 215-228, IRRI, Manila.

Li, C. C., 1970. Inheritance of the optimum and critical photoperiod in tropical rices. Bot. Bull. Acad. Sin. 11: 1-15.

Yamagata, H., Y. Okumoto and T. Tanisaka, 1986. Analysis of genes controlling heading time in Japanese rice. In Rice Genetics, p. 351-359. IRRI, Manila.

Yokoo, M. and H. Fujimaki, 1971. Tight linkage of blast-resistance with late maturity observed in different Indica varieties of rice. Jpn. J. Breed. 21: 35-39.

Yokoo, M. and F. Kikuchi, 1977. Multiple allelism of the locus controlling heading time in rice by the aid of the linkage with blast resistance. Jpn. J. Breed. 27: 123-130. (Japanese/English)

Yokoo, M., F. Kikuchi, A. Nakane and H. Fujimaki, 1980. Genetical analysis of heading data by aid of close linkage with blast resistance in rice. Bull. Nat. Inst. Agric. Sci. D31: 95-126.

Yu, C.J. and T. T. Yao, 1968. Genetische studien beim Reis. II. Die Koppelung des Langhullspelzengens mit dem Photoperiodizitatsgen. Bot. Bull. Acad. Sinica 9: 34-35.