ABSTRACT:Controlling combining chromosomes in meiosis is necessary to maintain genome constitutions. The Dryopteirs varia complex is one of the apogamous fern groups which perform recurrent hybridization with unequal meiosis, and produce spores after meiosis of the genome doubling in spore mother cells. We analyzed five low-copy nuclear markers (AK1, EST, GapCp, G6PD, and PgiC) and three regions of plastid DNA (rbL, ndhF and trnL-F). The genetic constitution of the five nuclear markers was the same in the D. varia complex. Therefore, recurrent hybridization seemed to be occurred without homoeologous chromosome paring.
KEYWORDS: Apogamous ferns, Homoeologous, Hybridization, Unequal meiosisControlling combining chromosomes in meiosis is necessary to maintain genome constitutions (Riley and Chapman 1958; Wall et al. 1971;Griffiths et al. 2006). Especially, meiosis of hybrids does not succeed because appropriate partners to pair are absent (Hirabayashi 1969(Hirabayashi , 1970. To solve this problem, allopolyploid has doubled set of chromosomes originated from progenitors to pair (Kihara 1924(Kihara , 1930 Wagner 1954; Riley and Chapman 1958;Ebihara et al. 2005;Griffiths et al. 2006; Sessa et al. 2012). Furthermore, allopolyploids can get more genome from other progenitors through recurrent hybridizations. In this case, the set of chromosomes originated from a parental allopolyploid and another parental diploid are also saved (Kihara 1924(Kihara , 1930 Riley and Chapman 1958;Griffiths et al. 2006; Sessa et al. 2012).Many apogamous ferns also have to control behaviors of their chromosomes because they produce spores through delicate mechanism called 'Döpp-Manton scheme' (Döpp 1939;Manton 1950). In this mechanism, disomic chromosome pairing and meiosis is occurred after genome doubling of spore mother cells. However, if homoeologous chromosomes pair randomly, offspring may have various morphological characteristics because genome constitutions of them are edited, but it does not seem to be occurred in regular shaped spores (Manton 1950). In fact, if homoeologous chromosome pairing is occurred in spore mother cells, the offspring get difficulty to survive (Otsuki et al. 2012).Otherwise, apogamous ferns get genetic diversity through hybridization between sexual species (Grusz et al. 2009;Ebihara et al. 2012;Dyer et al. 2012;Hori et al. 2014). Apogamous ferns get genetic variation through hybridization maintaining maturity of their spores in three patterns: (1) tetraploid apogamous hybrid between triploid apogamous species and diploid sexual species (Walker 1962; Watano and Iwatsuki 1988;Grusz et al. 2009;Dyer et al. 2012); (2) tetraploid apogamous hybrid between diploid apogamous species and tetraploid sexual species (Walker 1962); (3) triploid apogamous hybrid between diploid apogamous species and diploid sexual species (Walker 1962; Suzuki and Iwatsuki 1990;Chao et al. 2012;Jaruwattanaphan et al. 2013); (4) triploid apogamous hybrid between triploid apogamous species and diploid sexual species (Hori et al. 2014); (...