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Systematic Status of the Glycine Tomentella and G. Tabacina Species Complexes (Fabaceae) Based on ITS Sequences of Nuclear Ribosomal DNA

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Systematic Status of the Glycine Tomentella and G. Tabacina Species Complexes (Fabaceae) Based on ITS Sequences of Nuclear Ribosomal DNA J. Plant Res. 114: 435-442, 2001 Journal of Plant Research (C bv The Botanical Societv of Jaoan 1999 Systematic Status of the Glycine tomentella and G. tabacina Species Complexes (Fabaceae) Based on ITS Sequences of Nuclear Ribosomal DNA Yue-le C. Hsing’, Jaw-Shu Hsieh’, Ching-l Peng’, Chang-Hung Chou’ and Tzen-Yuh Chiang3* 1 Institute of Botany, Academia Sinica, Nankang, Taipei, Taiwan 115 Department of Agronomy, National Taiwan University, Taipei, Taiwan 107 3 Department of Biology, National Cheng-Kung University, Tainan, Taiwan 700 The phylogeny of the subgenus Glycine was reconstruct- studied extensively (Hymowitz and Newell 1978, Newell and ed based on nucleotide sequences of the internal tran- Hymowitz 1978,1980,Hymowitz and Singh 1987, Ohashi eta/. scribed spacer (ITS) region of the nuclear ribosomal DNA to 1991). Over the last two decades, the phylogeny of Glycine examine the systematic status of the G. fomentella and G. has also been elucidated based on isozyme data (Broue et fabacina species complexes. Rooted at the subgenus Soja a/. 1977), RFLP patterns of the chloroplast genome (Doyle et (2 species, 7 accessions), parsimony analysis was conduct- a/. 1990a),urease gene polymorphism (Menancio et a/. 1990), ed for 17 species (31 accessions) of the subgenus Glycine, and nuclear ribosomal DNA variation (Doyle and Beachy including 9 and 6 populations of G. fomentella s./. and G, 1985, Kollipara et a/. 1997, Singh et a/. 1998). tabacina s.l., respectively. The nrlTS phylogeny indicated Most Glycine species are diploid with a basic chromosome polyphyly of G. fomentella s.1. as well as G. fabacina s.l. In number of n=20 (Newell and Hymowitz 1983). Natural the G. fomentella species complex, larger legumes, narr- tetraploidy (2n=80) in G. tomentella, G. tabacina and G. ower leaflets, and deflexed indumentum hairs differentiated hirticaulis, and aneuploidy [2n=38,78 (fertile) and 79 (ster- G. dolichocarpa from G. fomentella S.S. The tetraploid G. ile)] in G. tomentella have been documented (Newell and dolichocarpa (2n= 80) and aneuploid G. fomentella (2n= Hymowitz 1983, Singh and Hymowitz 1985a,Singh et a/. 1988, 38) represented independent lineages from another clade of 1992, Tindale and Craven 1988,1993, Kollipara et a/. 1995). the remaining diploid (2n=40) and tetraploid species with a Genome types based on meiotic chromosome behavior were DD genome type. Tetraploid G. tabacina (2n=80) was designated and used for interpreting polyploidy (Singh and closely related to G. dolichocarpa instead of the diploid G. Hymowitz 1985; Kollipara et a/. 1997). Multiple origins of bbacina (2n=40) with a BB genome type. The nrlTS both G. tabacina and G. tomentella have been suggested phylogeny suggested allopolyploidy of G, dolichocarpa and (Grant et a/. 1984, Doyle and Brown 1989). Doyle et a/. of the tetraploid G. tabacina, both of which possibly share a (1990b) also suggested that there have been numerous common parental species with an AA genome type. Their origins involving reticulation in polyploid G. tabacina, based phylogenetic affinity also indicated biased inheritance of the on a phylogeny derived from the cpDNA polymorphism. nrDNA ITS and a possible dominant role of the AA genome. Multiple colonization events in Pacific islands were also Phylogenetically, G. dolichocarpa and allotetraploid G. suggested on the basis of presence of different plastome fabacina should be recognized as distinct species. types. Cytological and molecular studies of Kollipara et a/. (1994) indicated frequent allopolyploidization in the evolution Key words: Glycine - Glycine tabacina - Glycine of G. tomentella complex. A previous study based on RFLP tomeritella - nrDNA ITS - Phylogeny - Polyploidization patterns of seed proteins also detected polymorphism within G. tomentella, of which one strain with larger legumes shared two specific DNA fragments with G. tabacina (Hsing et a/. The genus Glycine Willd. consists of subgenera Glycine 1995). In contrast, the larger-legumed strain shared another and Soja, with 16 wild perennial species and two annual two DNA fragments with another G. fomentella that had species, respectively. The genus is distributed throughout legumes of a smaller size. Another investigation based on Australia, tropical Asia and East Asia (Newell and Hymowitz SDS-PAGE and Western blotting of seed proteins also 1980). Most species of the subgenus Glycine are geo- revealed genetic differentiation between short-pod, and graphically isolated from the subgenus Soja except for G. long-pod G. tomentella (Hsieh et a/. 2001). tomentella and G. tabacina (Newell and Hymowitz 1978, Not only did molecular evidence indicate genetic differen- Ohashi et a/. 1991). The systematics of Glycine has been tiation but morphological features were also diverse in the Glycine tomentella complex (Kollipara et a/. 1994). While * Corresponding author: [email protected] studying taxa of Taiwan, Ohashi et a/. (1991) observed 436 Y.C. Hsing et a/ morphological polymorphism in G. tomentella (sensu lato) were taken from Kollipara et a/. (1997), the information of and distinguished G. dolichocarpa from G. tomentella on the legume sizes was obtained from the taxonomic literature basis of its larger morphs. While fruits of the latter species (Table 1). had two to five seeds, legumes of G. dolichocarpa contained five to nine seeds. Upper calyx lobes that were united to 2/ DNA isolation, PCR, and nucleotide sequencing 3, and the dense, deflexed hairs on the petioles were also Seedlings of the above materials were ground in liquid distinct in G. dolichocarpa. The larger morphs of G. nitrogen and stored in a freezer at -70 C. Genomic DNA tomentella have been illustrated in earlier literature and was extracted from the powdered tissue according to the identified as G. tomentosa Benth. (Tang and Lin 1962, CTAB procedure (Doyle and Doyle 1987). PCR amplification Chuang and Huang 1966). Nevertheless, G. dolichocarpa was performed on a MJ PTC-100 Thermal Cycler using a pair has received little attention and has barely been mentioned of universal primers, ITS4 (5-TCCTCCGClTAlTGATATGC- in recent systematic molecular studies (e.g. Gu et a/. 1994, 3) and ITS18 (5-CGTAACAAGGTCCGTAGG-3) (O’Kane Kollipara et a/. 1997). The systematic status of G. doli- et a/. 1996). PCR was carried out in 100 pl reaction mixtures chocarpa and its phylogenetic relationship with other taxa still containing 10 ng of template DNA, 10 pI of 1OX reaction remain unclear. buffer, 10 pl of MgCk (25 mM), 10 pI of dNTP mix (8 mM), 10 The internal transcribed spacer (ITS) region of the pmole of each primer, 10 pI of 10% NP-40, and 4 U of Taq ribosomal DNA, one of the gene families in the nuclear polymerase (Promega, Madison, USA). PCR amplification genome, has been frequently used for phylogenetic study at conditions were 30 cycles of denaturation at 94 C for 45 s, inter- and infrageneric levels (Baldwin et a/.1995, Chiang and annealing at 49 C for 1 min 15 s, and extension at 72 C for 1 Schaal 2000) due to its homogeneity among units by con- min 15 s, followed by a final extension step at 72 C for 10 min certed evolution as well as fast evolutionary rates (Wojciech- and storage at 4C. PCR products were separated on an owski et a/. 1993; Li 1997). Kollipara et a/. (1997) and Singh agarose gel and bands were cut out and purified using a et a/. (1998) used the nucleotide sequences of the ITS region purification kit (Boehringer Mannheim, Mannheim, Germany). of nuclear ribosomal DNA to infer the phylogeny of the entire Eluted DNA fragments were directly sequenced. Cycle genus. Nevertheless, multiple samples from each species, sequencing with Taq polymerase was then performed by 32P including those from the Glycine tomentella and G. tabacina radioactive labeling using the fmol Sequencing System complex, were not included in the above phylogenetic (Promega). For sequencing, two additional primers ITS3 (5- analyses, in which mostly only a single individual represent- GCATCGATGAAGAACGTAGC-3) and ITS5.8 (5-ACTC- ing each taxon was sampled. GATGGTTCACGGGATT-3) were synthesized. Both In light of the above developments, this study was perfor- strands were sequenced from both ends, using the PCR med to examine the monophyly of Glycine tomentella sensu primers and the two new internal primers. lato, and the monophyly of the G. tabacina complex with diploid and tetraploid populations. We also examined the Sequence alignment and phylogenetic analyses genetic status of G. dolichocarpa and its relationship to the Nucleotide sequences were aligned with the program diploid G. tomentella. CLUSTAL V (Higgins et a/.1992). The fixed gap penalty was 35 and the floating penalty was 4. Cladistic analysis of Materials and Methods sequence data of the subgenus Glycine, rooted at the subgenus So&, was performed with Phylogenetic Analysis Plant materials Using Parsimony Program (PAUP, Version 3.1.1., Swofford Both Glycine tomentella s.l. and G. tabacina s.1. are wide- 1993). Heuristic searches were performed with TBR branch spread throughout Australia, tropical Asia, southern China, swapping, stepwise addition of 10 random replicates, ac- and Taiwan (Huang and Ohashi 1977). To examine the celerated transformation (ACCTRAN), an unconstrained monophyly of these species complexes, seven populations number of maximum trees, and retention of multiple most of G. tomentella sensu lato including both strains with large parsimonious trees (MULPARS). All characters were (G. dolichocarpa in this study, two populations) and small unweighted. Confidence values for clades were obtained legumes (G. tomentella s.s., five populations), and five popu- by bootstrapping (Felsenstein 1985) with 1000 replicates lations of G. tabacina complexes were sampled. Seeds (Hedges 1992). The nodes with bootstrap values greater were collected in natural habitats and germinated in the than 0.70, as a rule of thumb, are significantly supported greenhouse of Academia Sinica in Taipei under uniform with > 95% probability (Hillis and Bull 1993). A g7 test conditions (Table 1).
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