14Severe reproductive isolation (RI) exists between the two subspecies of rice, 15 Indica and Japonica, but in the wild ancestors no post-zygotic RI was found. The 16 studies about the establishment of the interspecies RI of rice are still rear. A pair of 17 rice hybrid sterility genes, DOPPELGANGER 1 (DPL1) and DOPPELGANGER 2 18 (DPL2), offers a convenient example to study the evolutionary history of RI genes. 19 Either of the two loci has one non-functional allele . The hybrid 20 pollen carrying both DPL1-and DPL2-will be sterility. 21 We collected 811 individuals: Oryza sativa (132), the two wild ancestors O. 22 nivara (296) and O. rufipogon (383) as well as 20 DPL1 and 34 DPL2 sequences of O. 23 sativa from on-line databases. We analysed the genetic and geographic pattern of 24DPLs in all three species to determine the origination regions of DPL1-and DPL2-. 25 The neutral test as well as the diversities of nucleotide and haplotype were used to 26 detect if selection shaped the pattern of DPLs. 27 We found that DPL1-and DPL2-of rice emerged from wild ancestor populations 28 in South Asia and South China through two respective domestications. Comparing 29 with the ancestral populations, DPL1-and DPL2-both showed reduce of diversities, 30 however their frequencies increased in rice. We assume that the reduce of diversities 31 due to the bottleneck effect of domestication while the loss of one copy was preferred 32 by artificial selection for cost savings.
33Reproductive isolation (RI) is considered a failure to transmit genetic materials 36 (Futuyma, 2013). Barriers that lead to RI are classified as pre-or postzygotic for 37 practical (Baack et al, 2015; Futuyma, 2013; Lafon-Placette et al, 2016; Rieseberg 38 and Willis, 2007; Widmer et al, 2009). Many genes related to RI have been identified 39 in various species (Fishman and Sweigart, 2018; Masly et al, 2006; Nadir et al, 2018; 40 Ouyang and Zhang, 2018; Zuellig and Sweigart, 2018), yet studies that aim to 41 elucidate the evolutionary histories of RI genes are still rare. 42 Rice (Oryza sativa) is an important crop as well as a model species. Strong 43 postzygotic RI was found between two subspecies of rice, Japonica and Indica. Many 44 loci answering for the hybrid incompatibility have been found (Ouyang et al, 2010; 45 Wang et al, 2014). Hundreds of loci were found related to hybrid incompatibility, 46 indicating that the incompatibility between Japonica and Indica has a complex 47 48 detected: S5 (Chen et al, 2008; Yang et al, 2012b), Sa (Long et al, 2008) and DPLs 49 (Mizuta et al, 2010). S5 leads to endoplasmic reticulum stress. Sa and DPLs both lead 50 to hybrid pollen sterility. On the contrary, the two wild ancestors of rice, O. nivara 51 and O. rufipogon, are compatible (Cai et al, 2019). Rice and the two wild ancestors 52 constitute a excellent system for studying the evolution history of RI genes.
53The DPL system offers a suitable example for exploring this phenomenon. The 54 DPL system contains only two genes, DOPPELGANGER 1 (DPL1)...