30. Further study of Xa-4b gene for resistance to bacterial blight of rice

Tsugufumi Ogawa 1,2, G.A. Busto1, R.E. Tabien 1 and G.S. Khush1

1) International Rice Research Institute, PO Box 933, Manila, Philippines

2) Present address: Tropical Agriculture Research Center, 1-2 Ohwashi, Tsukuba, Ibarakiken, 305 Japan


A single dominant gene for resistance to bacterial blight was identified in Semora Mangga by Librojo et al. (1976). When the allele test between this gene and Sz-4 was made, no susceptible plants were obtained. Thus this gene was considered allelic to Xa-4. However, cultivars with Xa-4 are resistant at maximum tillering and later stages but Semora Mangga shows resistance at booting stage. Therefore the dominant gene of Semora Mangga was designated as Xa-4b. The segregating populations were inoculated with race 1 of bacterial blight only. However, we now have 4 races of bacterial blight in the Philippines. We reevaluated the resistance of Semora Mangga with 4 Philippine races of bacterial blight and found it to be resistant to the four races. The cultivars with Xa-4 are resistant to race 1 only. We also crossed Semora Mangga with Java 14 which has Xa-3 for resistance. No susceptible plants were observed in the F\2\ of this cross. On the basis of this new data it was concluded that Semora Mangga has Xa-3 gene for resistance to bacterial blight - not Xa-4b (Ogawa et al. 1986). We have now analyzed many crosses between cultivars having Xa-3 on one hand and Xa-4 on the other and susceptible plants were rarely observed in the F\2\ populations. Thus Xa-3 and Xa-4 are closely linked. Therefore, non appearance of susceptible plants in the F\2\ of IR22 which has Xa-4 for resistance and Semora Mangga was due to very close linkage between Xa- 3 and Xa-4.

Sidhu et al. (1978, 1979) analyzed several rice cultivars genetically for resistance to bacterial blight using Philippine Race 1 only. The conclusion was that some cultivars had Xa-4 and others Xa-4b. We re-examined these cultivars by inoculating them with four Philippine races of bacterial blight. Most of the cultivars which were earlier supposed to have Xa-4b turned out to have Xa-3 for resistance (Table 1). Thus, like Semora Mangga, their genes were mistakenly designated as Xa-4b. Our recent investigations show that there are no cultivars with Xa-4b. Thus this gene symbol is redundant.



Table 1. Reaction of cultivars studied by Sidhu et al. (1979) to four Philippine races of bacterial blight

===============================================================================
Cultivar            IRRI           Reaction to Races             Estimated
                    Acc.      ==============================       gene
                    No.        1       2       3       4  
===============================================================================
Ketan Gondo puro   17859      41.4    33.0    39.4    32.5        None
Sintawati          18863       9.4    33.8    34.6    14.9        Xa-4
Bandja Ili         17216       5.1    32.4    34.4    21.4        Xa-4
Ketan Gondel       17857       1.3     0.5     0.5     0.7        Xa-3
Padi Tomat         18408       0.8     0.5     0.4     0.5        Xa-3
Plampeyan          18494       1.7     0.7     0.7     0.6        Xa-3
Rante              18608       8.6     9.3     6.9    13.9        Xa-3*
Remadja Bulu       18620       2.1     1.5     0.9     1.3        Xa-3
Rengesi            18628       4.9     1.3     8.1     1.3        Xa-3
Seksrek            18740       2.6     1.1     1.5     3.0        Xa-3
Slendoran          18902       0.9     4.0     5.0    12.5        Xa-5
Balacong           19232       4.8     1.2     3.6     4.3        Xa-3
Ketan Lumbu        16461       1.1     0.4     0.2     0.6        Xa-3
Ketan Gundel       17865       1.5     0.4     0.3     0.7        Xa-3
Ketan Temon        17967       0.6     0.6     2.5     0.4        Xa-3
Lenggang Genuk Bulu18119       1.4     0.8     0.4     0.4        Xa-3
Sampangan Perak    19677       5.5     1.5     1.1     1.9        Xa-3
Sampangan Kunig    18678       3.9     1.4     1.9     2.5        Xa-3
Pulu banra kaja    20073       0.7     0.8     0.4     0.5        Xa-3
Pulut Bongo        28397       1.0     0.5     0.3     0.2        Xa-3
Bulu Maggali       17348       1.9     0.4     1.4     0.9        Xa-3
Bulu Putih         17350      17.5    13.7    11.4    14.3        Xa-3
Camor              17366       0.8     0.5     0.5     0.3        Xa-3
Gaja Baru          17586       4.6    21.1    20.9    11.8        Xa-4
Gendjah Urang      17639       0.7     0.3     0.7     0.4        Xa-3
Gewal Bangaran     17651       2.7     0.8     0.8     1.0        Xa-3
Hegar Manah        17733       5.4    26.9     -      11.9        Xa-4*
Hegar Manah-2      17733       2.9     1.5    28.4     -          Xa-4+Xa-10
Ketan Gadjih       13512      19.7    12.6    16.6    20.0        ?*
Benong             13520       8.6     2.5     2.4     -          Xa-3
Walen              16685       4.5     1.4     1.6     2.6        Xa-3
Ketan Bendu        17838       3.4     0.9     1.0     0.7        Xa-3
Ketan Gobel        17851       6.8     3.8     4.2     2.3        Xa-3
Ketan Laler        17897       1.8     0.7     0.8     0.8        Xa-3
Ketan Pandan       17932       0.5     0.7     0.3     0.6        Xa-3
Ketan Saponjono    17959       0.5     0.5     0.4     0.5        Xa-3
Ketan Tjendoni     17970       2.9     0.7     1.0     1.0        Xa-3
Palotan Melati     18426       0.8     0.8     1.3     0.9        Xa-3
Parew Buwun        18456       2.3     1.0     0.7     1.6        Xa-3
Pare Djerah        18458      19.3    13.4    17.2    14.9        Xa-3
Keian Gundil       24568       0.4     0.6     0.5     0.6        Xa-3
Pulut Kemandi      24621       0.2     0.2     0.3     0.4        Xa-3
Remadja             6793       7.7    28.6    31.8    15.5        Xa-4*
Sidjangguk         18826       0.6     0.8     0.6     0.5        Xa-3
Item               14652       8.8    12.4     8.1    14.5        Xa-3
Andel              17153       1.0     1.0     0.7     0.7        Xa-3
Bajang             17183       7.4    40.7    39.8    15.7        Xa-4
Beak Ganggas       17253       2.0     1.1     0.8     1.0        Xa-3*
Bulu Djambu        17343       2.2     1.3     1.3     1.2        Xa-3
Djawalara          17471       1.8     1.0     1.1     1.9        Xa-3*
Djrabang           17543       1.5     0.9     0.8     1.2        Xa-3
Boder              17315       1.4     1.2     0.9     1.0        Xa-3
Bujtak Tuwunbatu   17356       3.8     1.8     1.4     1.8        Xa-3
Bulu Sampang       17352       1.4     1.9     1.2     1.4        Xa-3
Balik Semah        17201       2.7     2.5     1.4     2.2        Xa-3
Bulu Baselijan     17341       4.3     1.8     1.8     1.3        Xa-3
Indel              17748       4.5    21.9    28.0     7.6        Xa-3
===============================================================================
*segregation was observed
===============================================================================

From experience we have learned that resistance to bacterial blight should be evaluated on quantitative symptoms such as disease index, lesion area or lesion length. Resistant and susceptible plants are sometimes difficult to tell apart when scored at 14 days after inoculationl. Therefore we should pay more attention to the characteristics and variability of disease symptoms at 3-4 weeks after inoculation. Additional points to consider are:


1. The expression of lesion length is affected by sunshine and temperature during disease development, the genetic background of the cultivar, concentration of inoculation, and virulence.

2. Genetic analysis by inoculation at maximum tillering is less reliable as compared to the inoculation at booting stage.

3. It is desirable to evaluate the resistance by inoculating with several races.



References

Librojo, V., H.E. Kauffman and G.S. Khush, 1976. Genetic analysis of bacterial blight resistance in four varieties of rice. SABRAO J. 8: 105-110.

Ogawa, T., R.E. Tabien, G.A. Busto, G.S. Khush and T.W. Mew, 1986. The relationship of Xa-3, Xa-4 and Xa-4b for resistance to rice bacterial blight. RGN 3: 83-84.

Sidhu, G.S., G.S. Khush and T.W. Mew, 1978. Genetic analysis of bacterial blight resistance in seventy-four cultivars of rice, Oryza sativa L. Theor. Appl. Genet. 53: 105-111.

Sidhu, G.S., G.S. Khush and T.W. Mew, 1979. Genetic analysis of resistance to bacterial blight in seventy cultivar of rice, Oryza sativa L. from Indonesia. Crop Improv. 6: 19-25.