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Materials and methods

Plant materials
Forty-seven lines of alloplasmic or NC hybrids which have plasmons derived from various Triticum and Aegilops species combined with a single nuclear genome from Triticum aestivum L. em Thell. cv. Chinese Spring (CS) were used in this study. For some NC hybrids, their cytoplasms were replaced by that of CS after reverse crosses of the hybrids as male parents with CS as a female parent (see Results and discussion). The original hybrids were provided by Prof. K. Tsunewaki, Kyoto University (now Fukui Prefectural University). CS served as a euplasmic control.

RAPD-PCR detection of DNA polymorphisms
Total DNAs were extracted according to Liu et al. (1990) from one to five plants obtained after self-fertilization of fertile NC hybrids and from single progeny plants of male-sterile NC hybrids after backcrosses with the paternal CS. DNAs from the reverse F1 hybrids were also extracted in the same way. RAPD-PCR was performed in a reaction mixture (10 micro-liter) containing 10 ng of total DNA, 10 mM Tris-HCl (pH 8.3), 50 mM KCl, 2 mM MgCl2, 0.001% gelatin and 0.32 microM random decamer primer (Operon Technologies). PCR program was as follows: a pre-denaturation step for 30 s at 93C; and 40 cycles of 1 min at 93C, 1 min at 36C, 1.5 min at 72C; followed by postextension for 2 min at 72C. Amplified fragments were resolved by electrophoresis through 1.4 % agarose gels and visualized by staining with ethidium bromide.


Results and discussion

Polymorphic DNA fragments were detected by RAPD-PCR analysis using total DNAs extracted from 47 NC hybrid lines. Only 31 random primers were used in this study and seven were found to give polymorphic fragments among them. The polymorphisms detected were either amplification or non-amplification of DNA fragments unique in single lines or in several lines mostly with phylogenetically related cytoplasms.

Six among seven polymorphic fragments were assigned to the cytoplasmic genomes (Table 1). Three types of cytoplasmic polymorphisms were detected as amplification of single fragments. A primer OPA02 amplified a 1.6 kbp fragment (A021,600) in three NC hybrids with M plasmon of Ae. comosa thessalica (Kyoto University code number C05), Mh plasmon of Ae. herdreichii (C06) and N plasmon of Ae. uniaristata (C07) (Fig. 1A). The polymorphism in the male-sterile M and Mh plasmons was detected in the progeny plants obtained after backcrosses of the NC hybrids with the paternal CS and thus considered to be derived from the plasmons. The cytoplasmic origin of this polymorphism was confirmed by RAPD-PCR analysis using DNAs from the reverse hybrids in which their cytoplasms were replaced by that of CS. As expected the fragment A021,600 could not be amplified in the reverse hybrids (Fig. 2A). Another primer OPB19 amplified a 1.3 kbp fragments (B191,300) in the NC hybrids with Sb (C12, Ae. bicornis), T (C13, Ae. mutica) and T2 (C14, Ae. mutica) plasmons (Fig. 1B). OPE07 amplified a 3.2 kbp fragment (E073,200) in three NC hybrids with Sl (C10, Ae. sharonensis), Sb (C12, Ae. bicornis ) and Sv(C33, Ae. kotschyi ) but did not in NC hybrids with S (C08, C17, Ae. speltoides) and other S-derivative plasmons. They were assigned to the cytoplasmic variations in the same way using the reverse hybrids. The polymorphic fragment A021,600 detected in M and two M-related plasmons was absent in three NC hybrids with T and T2 plasmons of Ae. mutica and Mo plasmon of Ae. ovata. Mo plasmon of Ae. ovata is thought to have evolved from T plasmon of Ae. mutica after hybridization with Ae. umbellulata as a pollen parent followed by amphidiploidization (Tsunewaki 1995). Therefore, as far as this polymorphism is concerned, T plasmon of Ae.mutica can be considered to have remained unchanged in Mo, plasmon of Ae. ovata after its evolution. On the other hand, B191,300 was absent in a NC hybrid with Mo plasmon of Ae. ovata. This polymorphism thus can be considered to have evolved after amphidiploidization of Ae. ovata. A reason why this same polymorphism was present in Sb plasmon of remains unknown.

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