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Wide Somatic Hybrids of Citrus with Its Related Genera and Their Potential in Genetic Improvement

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Wide Somatic Hybrids of Citrus with Its Related Genera and Their Potential in Genetic Improvement Euphytica 118: 175–183, 2001. 175 © 2001 Kluwer Academic Publishers. Printed in the Netherlands. http://www.paper.edu.cn Wide somatic hybrids of Citrus with its related genera and their potential in genetic improvement W.W. Guo & X.X. Deng∗ National Key Laboratory of Crop Genetic Improvement, Huazhong Agricultural University, Wuhan 430070, China; (∗author for correspondence; e-mail: [email protected]) Received 11 November 1999; accepted 11 September 2000 Key words: Aurantioideae, Citrus, genetic improvement, protoplast fusion, sexually compatible / incompatible intergeneric somatic hybrids Summary Citrus wild relatives are an untapped germplasm reservoir, which possesses many elite resistance traits. Genetic introgression into Citrus by conventional methods has been greatly hindered by polyembryony, pollen / ovule sterility, sexual / graft incompatibility, long juvenility etc. Somatic hybridization via protoplast fusion may make it possible to transfer genes from wild relatives to Citrus. To date, more than sixty sexually compatible or incompat- ible intergeneric somatic hybrids between Citrus and its various related wild genera have been produced by PEG – or electrically – induced fusion. These wide somatic hybrids were identified by morphology, cytology, isozyme, RAPD and RFLP analyses. Genetic variation or recombination was also revealed in some of them. Several sexually compatible combinations have flowered and set fruits. The agronomic performance of these wide somatic hybrids is diverse, and the current results are not conclusive. Somatic hybrids are being tested as rootstocks. The prospects of wide somatic hybridization of Citrus with its wild relatives are discussed. Introduction resistance, and exhibits good cold hardiness (Bitters, 1969). Citropsis offers strong resistance to Phytoph- Citrus wild relatives are an untapped germplasm reser- thora induced diseases (Swingle & Reece, 1967) and voir, which possesses many elite resistance traits. Pon- burrowing nematode (Ford & Feder, 1960). Feronia cirus is highly resistant to citrus tristeza virus (CTV), is known to be drought tolerant, and due to its de- Phytophthora-induced diseases, as well as the citrus ciduous nature, it may be a source of genes for cold nematode (Castle, 1987), and is extensively used as a hardiness (Swingle & Reece, 1967). Swinglea glu- citrus rootstock in China and Japan. Fortunella is cold tinosa grows well in sandy loam and porous limestone hardy (Swingle & Reece, 1967), F. hindsii var. Chin- soils (Swingle & Reece, 1967), and Feroniella lucida tou was verified to be highly resistant to Phytophthora is dwarfing (Swingle & Reece, 1967), resistant to CTV parasitica (Zhu et al., 1991), and F. crassifolia cv. (Yoshida, 1996). Murraya paniculata is immune to Meiwa has been identified to be CTV resistant (Mestre citrus huanglongbing (Chen & Liao, 1982; Guo & et al., 1997). Microcitrus offers resistance to drought, Deng, 1998a), resistant to citrus nematodes (Sykes, flooding, the burrowing nematode (O’Bannon & Ford, 1987), citrus tristeza virus (Yoshida, 1996), and grows 1977), and Phytophthora induced diseases (Carpenter well in alkaline conditions (Sykes, 1987). Clausena is & Furr, 1962). Severinia offers cold resistance, salt tolerant to alkaline conditions (Sykes, 1987). Further and boron tolerance (Cooper, 1961), resistance to study of citrus wild relatives will undoubtedly reveal Phytophthora (Carpenter & Furr, 1962; Grimm & more elite traits. Hutchison, 1977), and nematodes (Baines et al., 1960; However, these citrus wild relatives have not been Hutchison & O’Bannon, 1972). Atalantia is known to commercially used except Poncirus, Fortunella,and perform well in wet soils, suggesting Phytophthora Clausena lansium, and only Poncirus and Fortun- ________________________________________________________________________________________中国科技论文在线176 http://www.paper.edu.cn ella are being successfully utilized through sexual ance of plants regenerated from most fusions between hybridization in breeding programs. Other citrus wild sexually incompatible parents was acceptable. Only a relatives performed poor as citrus rootstocks, or are few sexually incompatible fusions regenerated weak sexually / graft incompatible with citrus (Iwamasa et shoots (Motomura et al., 1996a; Shinozaki et al., 1992; al., 1988). Even if they are sexually compatible with Takayanagi et al., 1992). The success in regenera- citrus, the following problems or difficulties still exist tion of these somatic hybrids proved that protoplast (Barrett, 1985): 1) sexual crosses are usually much fusion was effective to bypass the difficulties such more difficult to make, and if seed is obtained at all, as polyembryony, pollen/ovule sterility, or sexual in- the seed yield is often low and it may be nonviable; compatibility encountered in sexual crossing, and it 2) if hybrids are obtained, the progenies ultimately is possible to realize genetic recombination of Citrus available for selection are often few in number due with its wild relatives. For the detailed advantages of to lethals, abnormal recombinants, or poor field sur- somatic hybridization versus sexual hybridization to vival; 3) the surviving hybrids may not flower, those recover hybrids among compatible and incompatible that do may have very long juvenile periods, or may parents, see Grosser & Gmitter, 1990a. be completely to partially ovule or pollen sterile; 4) undesirable traits may be integrated into Citrus.In comparison, somatic hybridization via protoplast fu- Genetic identification of these wide somatic sion is an alternative to circumvent some of these hybrids difficulties, and may pave the way for transferring their resistance genes to Citrus. These somatic hybrids were routinely identified by morphology, cytology, isozyme, RAPD (randomly amplified polymorphic DNA) (Grosser et al., 1996c; Somatic hybrids of Citrus with its related genera Shi et al., 1998b; Xiao et al., 1995) and RFLP (re- produced to date striction fragment length polymorphism) (Ohgawara et al., 1994; Motomura et al., 1996b), and genetic Mesophyll protoplasts isolated from citrus leaves can differences were revealed in some of them. not divide and regenerate into plants under the cur- rently used culture schemes (Tusa et al., 1992), but Leaf morphology protoplasts isolated from Citrus embryogenic callus can do so. Embryogenic calluses have been induced, The leaf of these wide somatic hybrids was generally and their protoplast-derived plants have been regener- wider and thicker than either fusion parent (Grosser ated from many species of Citrus genus and its related et al., 1996c) probably due to ploidy effect. Citrus genera (Huo et al., 1999; Jumin & Nito, 1996; Vardi & species are unifoliate. When citrus wild relatives with Spiegel-Roy, 1982). For protoplast fusion, the model compound leaves were used as the other fusion par- of ‘embryogenic callus + mesophyll protoplasts’ is ex- ent, their corresponding somatic hybrids were also of tensively used in Citrus, and allows the regeneration of compound leaves with bifoliate or trifoliate character, hybrid plants. For methods and procedures of Citrus as can be seen from the somatic hybrids of Citrus + protoplast isolation, protoplast fusion (PEG and elec- Poncirus (Ohgawara et al., 1985), Citrus + Citrop- trofusion) and culture, see Grosser & Gmitter, 1990a; sis (Grosser et al., 1996c), Citrus + Murraya (Guo Guo et al. 1998; Hidaka & Omura, 1992. & Deng, 1998b), Citrus + Clausena (Guo & Deng, Since the first intergeneric somatic hybrid in Citrus 1999b), and Citrus + Feronia (Grosser et al., 1996c). was regenerated (Ohgawara et al., 1985), more than Additionally, plants with unexpected abnormal leaf 60 intergeneric somatic hybrids have been obtained morphology were also found in some fusion combin- by PEG or electrically induced fusion (Table 1). Two ations (Louzada et al., 1993; Motomura et al., 1995, thirds of the existing wide somatic hybrids were cre- 1996b). ated by the research group led by Prof. J.W. Grosser in University of Florida, U.S.A., the rest produced in Ploidy variation France, Israel, Japan, and in our program led by Prof. X.X. Deng in China. From the published papers, it is Most wide somatic hybrids were tetraploids (2n = 4x = noteworthy that fusions between sexually compatible 36) following fusion of diploid + diploid, but un- parents regenerated healthy plants, and the perform- expected ploidy variation also exists in some fusion ________________________________________________________________________________________中国科技论文在线 http://www.paper.edu.cn177 Table 1. Somatic hybrids between Citrus and its related genera Embryogenic parent Mesophyll parent Number of fusion References combinations Sexually compatible Citrus aurantium Citrangea 2 Grosser et al., 1998, 2000 Poncirus 2 Grosser et al., 1994, 2000 Citrus deliciosa Fortunella japonica 1 Ollitrault et al., 1996 Poncirus 1 Ollitrault et al., 1996 C. jambhiri (hybrid Milam) Citrange 1 Grosser et al., 1998 Citrumelob 1 Grosser et al., 1998 Citrus paradisi Poncirus 3 Grosser et al., 1996c, 2000 Citrus reticulata Citrange 3 Grosser et al., 1996b, 1998; Guo, 1998 Citrumelo 1 Grosser et al., 1992 Fortunella obovata 1 Liu, 1999; Microcitrus papuana 1 Motomura et al., 1996b Poncirus 3 Grosser et al., 1992, 1994; Guo, 1998; Citrus sinensis Citrange 2 Louzada et al., 1992; Orgawara et al., 1991 Fortunella crassifolia 2 Deng et al., 1992; Grosser et al., 1996c Microcitrus papuana 2 Grosser et al., 1996c Poncirus 4 Grosser et al., 1988b, 1994, 2000; Orgawara
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