The Species of the Genus Oryza and Transfer of Useful Genes from Wild Species Into Cultivated Rice, O
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Breeding Science 60: 518–523 (2010) doi:10.1270/jsbbs.60.518 Review The species of the genus Oryza and transfer of useful genes from wild species into cultivated rice, O. sativa Kshirod K. Jena*1) 1) Plant Breeding, Genetics and Biotechnology Division, International Rice Research Institute, C/o National Institute of Crop Science, Rural Development Administration, Suwon 441-857, Republic of Korea The genus Oryza has 24 species out of which two are cultivated (O. sativa and O. glaberrima) and 22 are wild species. Of the 22 wild species, six are in the primary gene pool of O. sativa complex and these wild species are easily crossable with the major cultivated species. These have the same AA genome as O. sativa. However, there are 10 wild species under O. officinalis complex having BB, CC, BBCC, CCDD, EE and FF genomes. The wild species of this complex are in the secondary gene pool and are cross incompatible with O. sativa. There are six most distantly related wild species with either diploids or tetraploids of GG, HHJJ and HHKK genomes and are highly cross incompatible with O. sativa. All the 22 wild species of Oryza are a vast reservoir of genes for biotic and abiotic stresses resistance. Some of the yield enhancing traits/genes from AA genome wild species have been identified and mapped with molecular markers for their integration into O. sativa genome. A broad-spectrum resistance gene for bacterial blight resistance (Xa21) has been iden- tified in O. longistaminata and introduced into many rice cultivars. Advances in biotechnology have facili- tated the development of interspecific hybrids between O. sativa and wild species of secondary and tertiary gene pools. Some important genes Pi40 and Bph18 for resistance to blast and brown planthopper, respective- ly, have been successfully transferred into elite cultivars from O. australiensis and the function of one blast resistance gene (Pi9) derived from O. minuta is elucidated. Many important genes from the most distantly related wild species such as O. alta, O. granulata, O. longiglumis and O. coarctata are expected to be trans- ferred into cultivated rice in the future using the latest tools of molecular genetics and biotechnology. Key Words: Rice, wild species, genus Oryza, genes. Introduction ance to biotic and abiotic stresses and eventually increase yield potential of modern cultivars. The objective of this pa- Rice (Oryza sativa L.) is the most economically important per is to discuss the status of the species in the genus Oryza food crop in the world and provides two third of calorie in- and the transfer of high value genes present in wild species take of more than three billion people in Asia and one-third into rice cultivars using the tools of modern biotechnology of calorie intake of nearly 1.5 billion people in Africa and and genetics. Latin America (Khush 2005). Rice is cultivated worldwide under various agro-climatic conditions. However, rice pro- Wild species of Oryza duction in recent years has been affected seriously by major biotic and abiotic stresses due to adverse climatic change The genus Oryza of the Gramineae family has 24 species. and breakdown of resistance genes in elite cultivars Two of the 24 species, O. sativa L. and O. glaberrima (Normile 2008). The genetic variability for resistance or tol- Steud., are cultivated cereals and 22 are wild species distrib- erance to biotic stresses is limited in cultivated rice gene uted in different geographic locations worldwide (Khush pool but abundantly present in the gene pools of wild species 1997, Vaughan 1989). O. sativa is cultivated as a major ce- belonging to the genus Oryza. There is an urgent need to real crop in most parts of Asia and consumed as a staple broaden the gene pool of cultivated rice by transferring valu- food. The African cultivated rice, O. glaberrima is grown in able genes from wild species for enhancing resistance/toler- small areas in West Africa. O. sativa has two subspecies: japonica and indica. The subspecies japonica has a narrow Communicated by H. Yasui genetic resources compared to indica subspecies which has Received September 30, 2010. Accepted October 31, 2010. a wide genetic diversity. Oryza wild species were classified *Corresponding author (e-mail: [email protected]) into three main groups or complexes based on the ease of Genus Oryza and transfer of useful genes from wild species 519 gene transfer from wild species into cultivated rice. These gene transfer into cultivated rice difficult. are: (1) O. sativa complex, (2) O. officinalis complex. (3) There are two diploid wild species, O. granulata and O. meyeriana and O. ridleyi complex (Morishima and Oka O. meyeriana under O. meyeriana complex and possess the 1960). These groups/complexes were later named as prima- GG genome. These two species are cross incompatible with ry, secondary and tertiary gene pools of Oryza, respectively O. sativa. Two tetraploid (O. longiglumis and O. ridleyi) (Khush 1997). wild species with HHJJ genome on the other hand are in- The O. sativa complex has two cultivated species and six cluded in the O. ridleyi complex and these species are highly wild species with the AA genome (Table 1). These species cross-incompatible with the cultivated species, O. sativa. are diploid, cross compatible and show homologous chro- Two more wild species such as O. coarctata which was mosome pairing. The perennial wild species, O. rufipogon is previously called Porteresia coarctata and another species the progenitor of the cultivated Asian rice O. sativa while O. schlechteri are similarly included in the tertiary gene O. barthii is the progenitor of the cultivated African rice pools. These two species are tetraploid with the HHKK ge- O. glaberrima (Chang 1976, Oka 1988, Vaughan et al. nome (Ge et al. 1999, Khush 1997). 2008, Zhu and Ge 2005). Of the two Asian wild species, the perennial O. rufipogon is distributed throughout tropical Useful genes of wild species of Oryza Asia and Oceania, whereas the annual O. nivara is restricted to tropical continental Asia. The other wild species endemic The wild species of Oryza contains numerous genes of eco- to Africa, O. longistaminata is perennial and rhizomatous. nomic importance and are being used as alternate sources of Two other perennial wild species, O. meridionalis and resistance or tolerance to biotic and abiotic stresses to enrich O. glumaepatula are endemic to tropical Australia, and the cultivated rice gene pool (Table 2). The wild species of South and Central America, respectively. Many useful genes AA genome have useful genes such as resistance to grassy from these AA genome species have been transferred by stunt and tungro viruses and bacterial blight, source of cyto- interspecific hybridization and selection. plasmic male sterility for hybrid rice production, and resis- There are ten wild species in O. officinalis complex tance to flooding (Brar and Khush 1997). However, the wild which have a wide geographical distribution. The species are species belonging to secondary gene pool of Oryza are dis- either diploid or tetraploid with six different types of ge- tantly related to O. sativa. The wild species of this gene pool nomes: BB (O. punctata), CC (O. officinalis, O. rhizomatis have a wealth of valuable genes needed for rice improve- and O. eichingeri), BBCC (O. punctata and O. minuta), ment. These species have genes conferring resistance to CCDD (O. latifolia, O. alta and O. grandiglumis), EE brown planthopper, white-backed planthopper, green leaf- (O. australiensis) and FF (O. brachyantha). These species hopper, leaf and neck blast, bacterial leaf blight (BB), yellow are cross incompatible with the cultivated species, O. sativa stem borer, sheath blight and genes for adaptation to aerobic and show non-homologous chromosome pairing making soil, and high biomass production to increase yield potential Table 1. Wild species of Oryza with chromosome number, genome composition and their origin Wild species Chromosome Number Genome Origin O. rufipogon Griff. 24 AA Tropical Asia O. nivara Sharma et Shastry 24 AA Tropical Asia O. longistaminata Chev. et Roehr 24 AA Africa O. barthii Chev. et Roehr 24 AA Africa O. meridionalis Ng 24 AA Tropical Australia O. glumaepatula Steud. 24 AA South and Central America O. punctata Kotschy ex Steud. 24, 48 BB, BBCC Africa O. minuta J.S. Presl. ex C.B. Presl. 48 BBCC Philippines and Papua New Guinea O. officinalis Wall ex. Watt 24 CC Tropical Asia O. rhizomatis Vaughan 24 CC Sri Lanka O. eichingeri Peter 24 CC South Asia and East Africa O. latifolia Desv. 48 CCDD South America O. alta Swallen 48 CCDD South America O. grandiglumis Prod. 48 CCDD South America O. australiensis Domin. 24 EE Tropical Australia O. brachyantha Chev. et Roehr 24 FF Africa O. granulata Nees et Arn. ex. Watt 24 GG Southeast Asia O. meyeriana Baill 24 GG Southeast Asia O. longiglumis Jansen 48 HHJJ Indonesia O. ridleyi Hook 48 HHJJ South Asia O. schlechteri Pilger 24 HHKK Papua New Guinea O. coarctata Roxb. 48 HHKK India 520 Jena Table 2. Wild species of Oryza with useful traits Wild species Genome Useful traitsa O. rufipogon AA Source of CMS, stem elongation ability, resistance to BB, and tungro tolerance O. nivara AA Resistance to grassy stunt virus and BB O. longistaminata AA Resistance to BB O. meridionalis AA Stem elongation ability O. punctata BB, BBCC Resistance to BPH and ZLH O. minuta BBCC Resistance to sheath blight, blast, BB, BPH O. officinalis CC Resistance to BPH, WBPH and GLH O. eichingeri CC Resistance to BPH, WBPH and GLH O. latifolia CCDD Resistance to BPH, Higher biomass for yield O. alta CCDD Resistance to stem borer and high biomass O. grandiglumis CCDD Higher biomass for yield O. australiensis EE Resistance to BPH and blast O.