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Camel/A Sinensis L. (Cultivated Tea) and Its Wild Relatives Based on Randomly Amplified Polymorphic DNA and Organelle-Specific STS

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Camel/A Sinensis L. (Cultivated Tea) and Its Wild Relatives Based on Randomly Amplified Polymorphic DNA and Organelle-Specific STS Heredity 78(1997)603—611 Received 11 July 1996 An assessment of genetic diversity among Camel/a sinensis L. (cultivated tea) and its wild relatives based on randomly amplified polymorphic DNA and organelle-specific STS FRANCIS N. WACHIRA, WAYNE POWELL & ROBBIE WAUGH* Department of Cell and Mo/ecu/ar Genetics, Scottish Crop Research Institute, lnvergowrie, Dundee DD2 5DA, Scotland, U.K. Membersof the genus Camellia interbreed relatively freely and several natural species hybrids exist. Species introgression into the cultivated germplasm of tea, Camellia sinensis L. (0. Kuntz), from related Camellia species has been postulated, and it is thought that teas currently under cultivation are not archetypal varieties. Randomly amplified polymorphic DNAs (RAPDs) and organelle-specific polymerase chain reactions were used to establish the affin- ities among cultivated tea and its wild relatives. The measures of similarity obtained indicated that RAPDs were taxonomically informative in Came/ha, and the species relationships revealed were generally consistent with those obtained using morphological, compatibility and terpenoid affinities. Species-specific RAPD products and products potentially diagnostic of introgressive hybridization into the cultivated gene pool were identified. The organellar genomes were remarkably conserved, with polymorphism detected in only one of four noncoding regions in the chioroplast and mitochondrial genomes. Keywords: Camelliaspp., gene introgression,phenetics, randomly amplified polymorphic DNA, similarity. Introduction several, including C. taliensis, C. grandibractiata, C. kwangsiensis, C. gymnogyna, C. crassicolumna, C. Afundamental goal of germplasm collection and tachangensis, C. ptilophylla and C. irrawadiensis, are conservation is the understanding of genetic rela- already used in parts of Asia (Chang & Bartholo- tionships within and between the species of concern. mew, 1984). In addition, seed from C. japonica and A good understanding is critical for the effective C. oleifera have been widely used for oil extraction organization and management of such germplasm in Japan and China, respectively (Sealy, 1958), and collections. In plant breeding programmes, estimates most Camelhia spp. are of great ornamental value. of genetic relationships can be useful for the identi- During the past century, a number of studies have fication of parents for hybridization, and for reduc- examined the relationships between Camellia species ing the number of accessions needed to maintain a in cultivation (Sealy, 1958; Chang & Bartholomew, broad range of genetic variability. The genus Camel- 1984 and references cited therein; Chuangxing, 1988; ha is composed of over 80 taxa (Sealy, 1958), of Tien-Lu, 1992). In general, morphological charac- which only one, C. sinensis L. (0. Kuntz), is ters, phytochemicals and terpenoids have been used. currently used commercially as a source of the However, it has been argued that these may not beverage tea. The potential for economic use of reflect the true level of genetic differentiation, as other species as a beverage is, however, real, and most are subject to large environmental effects. Leaf terpenoids have, nevertheless, proved useful in analysing closely related species (Takeo, 1983; *Corresponden. E-mail: [email protected] Nagata, 1986; Owuor et a!., 1987; McDowell et al., tPermanent address: Tea Research Foundation of Kenya, P0 1995), but at higher levels of classification they are Box 820, Kericho, Kenya. considered to be less useful owing to convergence 1997 The Genetical Society of Great Britain. 603 604 F. N. WACHIRA ETAL. and reticulate evolution. Ackerman (1973) and species is, therefore, vital for the efficient selection Takeda (1990) have evaluated cross-compatibility of of parents for interspecific hybridization. several Camellia species to determine their phenetic Molecular markers, such as the restriction frag- relationships. Because most species within the genus ment length polymorphisms (RFLPs) and randomly hybridize freely, several species complexes have amplified polymorphic DNAs (RAPDs), provide an been formed that have ultimately made evaluation efficient means of estimating genetic relationships of relationships even more daunting. Several minor (Ecke & Michaelis, 1990; Miller & Tanksley, 1990; taxa have been treated as conspecific with major Jung et at., 1993). Because of their simplicity, taxa, although more recently accumulated evidence RAPDs have been widely used in establishing rela- has shown that these minor taxa have no natural tionships within and between different species distribution and are derived from hybridization (Adams & Demeke, 1993; Ratnaparkhe et at., 1995; events involving different species (Parks et al., 1967; Yamagishi, 1995; Hoey et at., 1996; Orozco-Castillo Uemoto et a!., 1980). By analysing for the presence et at., 1996). In Camellia sinensis, RAPDs have been and distribution of phytochemicals (e.g. eugenol demonstrated to be useful in genotype differentia- glycoside, sasanquin and fluorescent flavonoid tion within cultivated germplasm (Wachira et at., sulphates) in a wide range of Camel/ia species and 1995). Variation in chloroplast (cp) and mitochon- their hybrids, Parks et a!. (1981) were able to drial (mt) DNAs has been used to examine relation- demonstrate the introgression of C. japonica traits ships among distantly related taxa (Waugh et a!., into C. sasan qua giving hybrids that were treated as 1990; van de Ven et at., 1993; Olmstead & Palmer, a distinct species (C. hiematis) by Chang (1981). 1994). These studies have been simplified by the Two of the major monographers of the genus, design of consensus primers that can be used in Sealy (1958) and Chang (Chang & Bartholomew, PCR to amplify homologous regions in different 1984), have identified two patterns of species rela- species and have proved to be highly informative tionships with very different arrangements of subge- (Taberlet et a!., 1991; Demesure et a!., 1995). Here, nera and generic sections. The more recent of these we report the use of RAPD markers and PCR studies (Chang & Bartholomew, 1984) has recog- amplification of chloroplast and mitochondrial nized Camellia as comprising two interfertile sequences to assess relationships among cultivated ancestor groups. Ancestor group 1 comprises one tea and allied wild species of the genus Cametlia. distinct subgeneric group, Camellia, and a second The results obtained have been compared with exist- ill-defined complex group, Protocamellia. Ancestor ing information based on cross-compatibility and group 2 is represented by two distinct subgenera, morphological markers used mostly in present-day Thea and Metacamellia. The subgenus Protocamellia taxonomy. has morphological characteristics of all the other three subgenera but does not merit treatment as a Materials and methods distinct genus (Chang & Bartholomew, 1984). The subgenus Thea has eight sections, one of which Plant mater/al (section Thea) comprises cultivated tea (Chang & Twenty Camellia species belonging to eight sections Bartholomew, 1984; Chuangxing, 1988). Tien-Lu of the genus Camellia were used in this study (Table (1992), however, recognized only seven sections in 1). These included nine tea cultivars (C. sinensis) the subgenus Thea. It has been proposed that all and two closely related wild species, C. irrawadiensis Camellia have a common ancestor and evolved from and C. tatiensis. A total of 28 genotypes were the primitive Protocamellia in two distinct directions, evaluated. one for Thea and Metacamellia and the other for Camellia (Chang & Bartholomew, 1984). DNA Species compatibility studies within the genus isolation and PCR ampilfication •have shown that most interspecific (inter- and intra- DNAwas isolated using the modified method of sectional as well as intrageneric) crosses can be Gawel & Jarret (1991) as described by Orozco- made without much difficulty, although some groups Castillo et a!. (1994). RAPD reactions were conduc- are more compatible than others (Ackerman, 1973; ted on individual DNA samples as described by Takeda, 1990). This provides opportunities for Wachira et al. (1995). Numerous adjustments were breeders to exploit the wide germplasm of Camel/ia initially examined in order to optimize the RAPD by looking at other species for traits of interest in assay to achieve the necessary reproducibility and their breeding endeavours. Knowledge of genetic resolution for all Camettia spp. The arbitrary relationships between tea and the other related sequence lO-mer primers used were either obtained The Genetical Society of Great Britain, Heredity, 78, 603—611. RELATIONSHIPS WITHIN THE GENUS CAMELLIA 605 Table 1 Camellia genotypes studied and their taxonomic affinities* Subgenus Section Species Source Camel/ia Came/ha C. pitardii NRIVOT, Japan C. saluensis NRIVOT, Japan Paracamelhia C. brevisyla NRIVOT, Japan C. kissi NRIVOT, Japan C. miyagii NRIVOT, Japan C. tenuifiora NRIVOT, Japan Oleifera C. oleifera NRIVOT, Japan C. susan qua NRIVOT, Japan Furfuracea C. furfuracea NRIVOT, Japan Protocamellia Arc hecamelhia C. granthamiana NRIVOT, Japan Metacamellia Camehliopsis C. assimilis NRIVOT, Japan C. sahicifolia NRIVOT, Japan Theopsis C. fraterna NRIVOT, Japan C. lutchuensis NRIVOT, Japan C, nokoensis NRIVOT, Japan C. rosaeflora NRIVOT, Japan C. tsai NRIVOT, Japan Thea Thea C. irrawadiensis TRFK, Kenya C. tahiensis NRIVOT, Japan C. sinensis(var. sinensis) Yabukita NRIVOT, Japan C. sinensis (var. assamica) AK1296 NRIVOT, Japan C. sinensis
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