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      Fine Scale Genomic Signals of Admixture and Alien Introgression among Asian Rice Landraces

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          Abstract

          Modern rice cultivars are adapted to a range of environmental conditions and human preferences. At the root of this diversity is a marked genetic structure, owing to multiple foundation events. Admixture and recurrent introgression from wild sources have played upon this base to produce the myriad adaptations existing today. Genome-wide studies bring support to this idea, but understanding the history and nature of particular genetic adaptations requires the identification of specific patterns of genetic exchange. In this study, we explore the patterns of haplotype similarity along the genomes of a subset of rice cultivars available in the 3,000 Rice Genomes data set. We begin by establishing a custom method of classification based on a combination of dimensionality reduction and kernel density estimation. Through simulations, the behavior of this classifier is studied under scenarios of varying genetic divergence, admixture, and alien introgression. Finally, the method is applied to local haplotypes along the genome of a Core set of Asian Landraces. Taking the Japonica, Indica, and cAus groups as references, we find evidence of reciprocal introgressions covering 2.6% of reference genomes on average. Structured signals of introgression among reference accessions are discussed. We extend the analysis to elucidate the genetic structure of the group circum-Basmati: we delimit regions of Japonica, cAus, and Indica origin, as well as regions outlier to these groups (13% on average). Finally, the approach used highlights regions of partial to complete loss of structure that can be attributed to selective pressures during domestication.

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          Interpreting principal component analyses of spatial population genetic variation.

          Nearly 30 years ago, Cavalli-Sforza et al. pioneered the use of principal component analysis (PCA) in population genetics and used PCA to produce maps summarizing human genetic variation across continental regions. They interpreted gradient and wave patterns in these maps as signatures of specific migration events. These interpretations have been controversial, but influential, and the use of PCA has become widespread in analysis of population genetics data. However, the behavior of PCA for genetic data showing continuous spatial variation, such as might exist within human continental groups, has been less well characterized. Here, we find that gradients and waves observed in Cavalli-Sforza et al.'s maps resemble sinusoidal mathematical artifacts that arise generally when PCA is applied to spatial data, implying that the patterns do not necessarily reflect specific migration events. Our findings aid interpretation of PCA results and suggest how PCA can help correct for continuous population structure in association studies.
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            Origin, dispersal, cultivation and variation of rice.

            There are two cultivated and twenty-one wild species of genus Oryza. O. sativa, the Asian cultivated rice is grown all over the world. The African cultivated rice, O. glaberrima is grown on a small scale in West Africa. The genus Oryza probably originated about 130 million years ago in Gondwanaland and different species got distributed into different continents with the breakup of Gondwanaland. The cultivated species originated from a common ancestor with AA genome. Perennial and annual ancestors of O. sativa are O. rufipogon and O. nivara and those of O. glaberrima are O. longistaminata, O. breviligulata and O. glaberrima probably domesticated in Niger river delta. Varieties of O. sativa are classified into six groups on the basis of genetic affinity. Widely known indica rices correspond to group I and japonicas to group VI. The so called javanica rices also belong to group VI and are designated as tropical japonicas in contrast to temperate japonicas grown in temperate climate. Indica and japonica rices had a polyphyletic origin. Indicas were probably domesticated in the foothills of Himalayas in Eastern India and japonicas somewhere in South China. The indica rices dispersed throughout the tropics and subtropics from India. The japonica rices moved northward from South China and became the temperate ecotype. They also moved southward to Southeast Asia and from there to West Africa and Brazil and became tropical ecotype. Rice is now grown between 55 degrees N and 36 degrees S latitudes. It is grown under diverse growing conditions such as irrigated, rainfed lowland, rainfed upland and floodprone ecosystems. Human selection and adaptation to diverse environments has resulted in numerous cultivars. It is estimated that about 120,000 varieties of rice exist in the world. After the establishment of International Rice Research Institute in 1960, rice varietal improvement was intensified and high yielding varieties were developed. These varieties are now planted to 70% of world's riceland. Rice production doubled between 1966 and 1990 due to large scale adoption of these improved varieties. Rice production must increase by 60% by 2025 to feed the additional rice consumers. New tools of molecular and cellular biology such as anther culture, molecular marker aided selection and genetic engineering will play increasing role in rice improvement.
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              Isozymes and classification of Asian rice varieties.

              Enzyme variation detected by starch gel electrophoresis was used to investigate the genetic structure of Oryza sativa L. species. Fifteen polymorphic loci coding for 8 enzymes were surveyed among 1688 traditional rices from Asia. Multivariate analysis of the data resulted in identification of six varietal groups, with two major ones, groups I and VI, two minor ones, groups II and V, and two satellite ones, groups III and IV. Group I is found throughout tropical Asia; it encompasses most Aman rices in Bangladesh, the Tjereh rices in Indonesia and the Hsien rices in China. Group VI is found mostly in temperate regions and in high elevation areas in the tropics; it encompasses most upland rices from Southeast Asia, the Bulu rices from Indonesia and the Keng rices from China. Groups II, III, IV and V share common differences from groups I and VI which suggest an alternative evolutionary history. Groups II and V are found in the Indian subcontinent from Iran to Burma. Well-known components of these are Aus rices from Bangladesh for group II and Basmati rices from Pakistan and India for group V. Groups III and IV are restricted to some deepwater rices in Bangladesh and Northeast India. Based on analogy with other classifications, Group I might be considered as the "Indica" type and Group VI as the "Japonica" type. Such terms, however, have a depreciated meaning due to discrepancies among various classifications.
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                Author and article information

                Contributors
                Role: Associate Editor
                Journal
                Genome Biol Evol
                Genome Biol Evol
                gbe
                Genome Biology and Evolution
                Oxford University Press
                1759-6653
                May 2019
                16 April 2019
                16 April 2019
                : 11
                : 5
                : 1358-1373
                Affiliations
                [1 ]UMR AGAP, CIRAD, Montpellier, France
                [2 ]UMR AGAP, Université de Montpellier, France
                [3 ]International Rice Research Institute (IRRI), Los Baños, Philippines
                Author notes
                Corresponding author: E-mail: glaszmann@ 123456cirad.fr .
                Author information
                http://orcid.org/0000-0002-4548-4606
                Article
                evz084
                10.1093/gbe/evz084
                6499253
                31002105
                50204337-f86b-4a24-a69d-7adee888c871
                © The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution.

                This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License ( http://creativecommons.org/licenses/by-nc/4.0/), which permits non-commercial re-use, distribution, and reproduction in any medium, provided the original work is properly cited. For commercial re-use, please contact journals.permissions@oup.com

                History
                : 11 April 2019
                Page count
                Pages: 16
                Funding
                Funded by: CIRAD 10.13039/501100007204
                Award ID: 1504-006
                Funded by: Labex Agro
                Award ID: ANR-10-LABX-0001-01
                Funded by: Investissements d'Avenir
                Award ID: ANR-16-IDEX-0006
                Categories
                Research Article

                Genetics
                3,000 rice genomes,oryza sativa,snps,local haplotypes,population structure,kernel density estimation

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