中国花生地方品种与育成品种的遗传多样性

以中国花生小核心种质中涉及来源于中国本土的145份地方品种和67份育成品种为材料,应用SSR技术从206对SSR引物中筛选出25对扩增效果好的多态性引物进行检测,并对其进行遗传多样性分析比较.结果表明:(1)地方品种与育成品种各具有特殊带型及各自独特的遗传特性.相似系数和多态性信息量均表明,地方品种的多样性比育成品种丰富,其中:地方品种之间的相似系数为0.57～0.99,平均0.795,多态性信息量0.530 0;育成品种之间的相似系数0.63～0.99,平均0.810,多态性信息量0.463 3.地方品种与育成品种之间的平均相似系数为0.794,变异范围0.56～0.99.(2)对不同生态区来源的分析表明,除黄河流域外其他各生态区地方品种的观测等位基因数和遗传多样性指数(分别为2.740 7～3.518 5和0.816 4～0.879 4)均比育成品种的对应值(分别为1.7012～2.145 6和0.4829～0.802 2)大,并以长江流域生态区地方品种的观测等位基因数和遗传多样性指数最大,分别为3.518 5和0.879 4.(3)聚类分析结果表明,花生核心种质中,中国本土资源分为3个品种群,即地方品种密枝亚种群、地方品种疏枝亚种群和育成品种群,与花生的亚种分类一致.(4)通过遗传多样性分析,鉴定出一批遗传差异较大的材料,其中zhh1398与zhh0041的遗传差异最大,相似系数为0.56,为花生品种的遗传改良及作图群体的构建奠定了基础.     

东北地区谷子地方品种和育成品种表型比较分析

我国东北地区谷子育种经历了从地方品种到育成品种的转变,但其具体表型性状等变化尚不清楚。本研究在敖汉旗对我国东北地区120个谷子地方品种和育成品种进行表型鉴定,并分析了遗传多样性,结果表明:东北地区谷子品种以绿幼苗、绿叶鞘、纺锤松散穗、黄粒、黄米型为主。育成品种晚熟、生物量大且高产,表现在出苗至抽穗、开花、成熟的时间以及播种至成熟的时间均极显著大于地方品种;主茎直径、叶片数、旗叶宽度均极显著大于地方品种;主穗重、主穗粒数极显著大于地方品种,主穗粒重和产量显著大于地方品种,但千粒重极显著小于地方品种,说明育成品种主要通过增加穗粒数来实现产量提升。遗传多样性指数分析发现,参试品种各性状遗传多样性指数大小顺序为:产量性状(2.0063)形态性状(1.9655)生育期(1.7238)质量性状(0.6370);地方品种护颖颜色、粒色、米色遗传多样性指数明显高于育成品种,说明谷子育种使粒色米色越来越趋于一致。基于30个性状的标准化数据,将参试品种聚为两类,聚类结果与品种来源地没有明显关联。类群Ⅰ为早熟、低产品种,且幼苗叶色、米色均为绿色和黄色;类群Ⅱ为晚熟、高产品种。根据10个主要表型性状,筛选出综合性状优良的红谷子、燕谷18等种质资源,可供东北地区谷子育种重点利用。     

SSR分子标记检测出的花生类型内遗传变异

花生是我国重要的食用油和蛋白质来源作物,鉴定其DNA分子多态性对品种改良和资源评价具有重要的意义。从已公布的花生Genomic-SSR和EST-SSR引物中筛选出34对引物,用来分别鉴定花生4大类型各24份共96份品种资源的分子变异,其中龙生型资源全部来自广西,普通型资源中有11份从国外引进,有13份来自广西和国内其他省市,多粒型资源只有两份来自中国,其他22份分别来自印度、美国和非洲等地,珍珠豆型资源中有22份是来自中国各地的育成品种或农家品种,有2份来自国外。研究结果为:分别有10～16对SSR引物能在4大类型花生资源中扩增出多态性DNA片段;这些多态性SSR引物都具有多位点特性;首次为SSR分子标记设立了一个新的评价指标——区别指数,多态性SSR引物的区别指数最高达0.992;资源间的平均遗传距离,多粒型为0.59,普通型为0.48,珍珠豆型为0.38,龙生型为0.17。根据遗传距离采用最长距离法对4大类型花生资源分别进行了聚类分析,构建了资源间的遗传关系图,花生4大类型可进一步分成不同类群,资源间的亲缘关系与其来源相关。观察到PM15和PMc297的扩增产物具有类型特异性,PM15能在龙生型、普通型和多粒型花生资源中扩增出多态性条带,而在珍珠豆型花生中扩增条带完全相同,PMc297也有相似的扩增结果。由于在多粒型花生资源中检测出的遗传多样性最丰富,研究结果支持西班牙专家Krapovickas 1994年公布的花生栽培种分类系统。总之在花生4大类型内资源中能检测出丰富的SSR分子标记,开发出更多的SSR分子标记将能充分揭示花生分子水平的变异,从而使花生遗传图谱构建、分子标记辅助育种成为可能。     

太湖流域粳稻地方品种遗传多样性研究

太湖流域稻作历史悠久,是我国粳稻高产地区之一。但育成品种的遗传基础日趋狭窄,将会给农业生产带来潜在风险。为了有效利用地方水稻资源以扩大改良品种的遗传基础,作者通过调查19个农艺性状,对823个太湖流域粳稻地方品种的遗传多样性进行了研究。结果表明:15个数量性状的变异系数(CV)范围在4.6%(结实率)和33.7%(单株产量)之间。颖壳色、稃尖色、芒型和粒型4个质量性状分别有10、6、5、4种表现型。19个农艺性状的多样性指数(H′)变化范围在0.757(粒型)和1.930(每穗实粒数)之间,平均为1.540。9月10日以后抽穗的品种比9月10日及其以前抽穗的品种具有更丰富的表型变异。主成分分析表明,株高、单株有效穗、每穗总粒数、每穗实粒数、着粒密度、千粒重、单株产量和穗颈长等8个性状是解释太湖流域粳稻地方品种多样性的重要性状。     

甘、青两省春小麦遗传多样性演变

对甘、青两省自20世纪40年代以来生产上曾大面积应用的200多个春小麦地方品种和育成品种资料进行了分析,结果表明,19个形态性状,5个农艺性状和2个品质性状都存在着广泛的遗传多样性,虽然育成品种农艺性状和品质性状的遗传多样性总体上大于地方品种,但是从20世纪60年代开展春小麦杂交育种工作之后,育成品种形态性状,农艺性状和品质性状遗传多样性呈下降趋势,评价育种对小麦遗传多样性的影响,不能足将地方品种和育成品种的遗传多样性作一简单比较就得出结论,而是应该在遗传多样性演变的动态过程中去研究。     

应用SSR标记分析大豆种质资源的遗传多样性

利用SSR分子标记分析了119个大豆品种的遗传多样性,结果表明：30对SSR引物在119份材料中共检测出159个等位变异,平均每对引物检测到5．30个等位变异;河北省农家品种中平均每对引物检测到5．17个等住变异,育成品种4．87个,省外品种4．93个,表明地方品种的遗传多样性高于育成品种。河北省农家品种、育成品种和省外育成品种依据SSR数据获得的品种间相似系数总体平均值相近,分别为0．698、0．698、0．672,但河北省农家品种较育成品种具有较大的变化幅度。119个品种可被划分为3个类群,在一定程度上能把育成品种和农家品种分开,并反映了一定的品种地域来源。     

应用Weitzman方法分析华中型猪品种的遗传多样性

本文根据华中型20个猪品种27个微卫星DNA标记的研究数据, 应用Weitzman方法, 即通过估计总体遗传多样性、期望多样性、品种对总体遗传多样性的贡献、边际遗传多样性、保种潜力等指标, 评估华中型20个猪品种的遗传多样性, 并通过构建品种间遗传多样性的最大似然树, 图示化展示20个品种间遗传多样性的相互关系。20个华中型猪品种的总体遗传多样性是11,707, 期望多样性占总的遗传多样性的66.96%, 金华猪、皖南花猪、嵊县花猪和乐平猪是对总体遗传多样性贡献最大的4个品种, 其贡献率分别是8.90%、7.46%、7.40%和7.04%, 保种潜力最大的4个品种分别是金华猪、嵊县花猪、杭猪和大花白猪。根据遗传多样性进行聚类分析, 大致可将华中型猪分为3类: 大围子猪、沙子岭猪、宁乡猪等分布于湖南、湖北的品种为一类; 南城猪、嵊县花猪与杭猪聚成另一类; 金华猪与皖南花猪单独聚成一类。研究结果可为我国华中型地方猪种遗传多样性的最大化保护和利用提供科学决策依据, 本文也对Weitzman方法在应用中的关键性问题进行了探讨。     

宁夏不同年代水稻品种的遗传多样性比较

对宁夏不同阶段主栽水稻品种(系)的主要农艺性状比较表明,第1阶段(1950-1962年)宁夏地方品种与第2阶段(1978-1989年)、第3阶段(1990-1999年)、第4阶段(2000-2005年)、第5阶段(2006-2012年)、第6阶段(2010-2013年)育成品种(系)的农艺性状差异显著,说明通过育种手段宁夏水稻品种的主要农艺性状得到了改良;从第2阶段到第6阶段的变化趋势为生育天数变长,有效穗数减少,穗长、穗粒数、单株粒重、单穗粒重、产量等增加,说明产量性状的育种改良从穗数型向穗粒兼顾型或穗重型发展。利用48对SSR引物对76份宁夏不同年代水稻品种(系)进行遗传多样性分析表明,共检测出290个等位基因,每个位点等位基因变幅为3~15个,平均6.04个;稀有等位基因有109个,占等位基因总数的37.59%;多态信息含量(PIC)变异范围为0.1423~0.8783,平均为0.5512;Nei’s基因多样性指数(He)的变幅为0.1492~0.8945,平均为0.6032。RM333、RM297、RM249、RM501和RM206引物表现为较高的等位基因数和稀有等位基因数,认为较适合应用于宁夏水稻品种的遗传多样性检测。分子方差分析(AMOVA)结果表明,6个不同阶段间遗传变异占总体变异的19.27%,各阶段内品种间的遗传变异占总体变异的80.73%。参试水稻UPGMA聚类分析表明,在遗传相似系数(GS)为0.805处可将其划分为5大类群,12个地方品种都被聚为第Ⅰ类群,具有外来引进血缘的8个常规粳稻品种被聚为第Ⅱ类群,5个优质常规粳稻被聚为第Ⅲ类群,第Ⅴ类群包括2个大穗高产品种,49个宁夏常规粳稻选育品种被聚为第Ⅳ类,说明大多数宁夏水稻品种的亲缘关系较近。6个不同阶段品种聚类分析,第1阶段(地方品种)独聚一类,第2、第3、第4阶段与第5、第6阶段育成品种亲缘关系相对较远;表明不同年代宁夏水稻遗传多样性有较大差异,2005年后宁夏水稻品种的遗传多样性趋于变小。宁夏水稻育种中应加强国内外种质资源的引进和利用,深入挖掘宁夏稻区地方品种和杂草稻资源的有利基因,拓宽宁夏水稻品种的遗传基础,以促进宁夏水稻育种的快速发展。     

基于DUS测试的上海粳稻地方品种遗传多样性分析

利用水稻DUS测试指南(2012版)中的38个水稻标准品种和52个基本测试性状对上海地区的172个粳稻地方品种进行遗传多样性分析,并与14个上海地区选育的申请品种保护的粳稻品种作比较。结果表明:地方品种表型性状共检测到176个等位变异,平均每个测试性状检测到3.6667个,变幅介于1~8个。Shannon's多样性指数(H')平均值为0.6834,变幅介于0~1.9327,77.08%的性状存在3个及以上的等位变异;地方品种的等位变异数、每个性状的等位变异数及其变异幅度、多样性指数平均值等均大于现代育成品种,在外观形态上,地方品种的遗传多样性比现代育成品种更丰富。利用UPGMA法进行聚类分析发现在相似系数为0.53时,可将186个粳稻品种分为3个类(亚)群,现代育成品种单独组成一个亚群;而主坐标分析将186个粳稻品种明显的分为地方品种和现代育成品种两大区。地方品种具有丰富的多样性,与现代育成品种具有明显的形态差异,可为今后粳稻育成品种的改良提供更多的形态基础。     

花生表型及SSR遗传多样性的研究

对山西省农业科学院经济作物研究所保存的75份花生材料(包括28个已审定的花生品种和47个地方品种)进行了包括株型、茸毛的有无、叶色、粒形、叶形、生长习性、开花习性、粒大小、粒色等表型性状的Shannon-Weaver遗传多样性指数(简称SWI)和Simpson遗传多样性指数(简称SI)分析。结果表明:参试的75份花生品种遗传多样性指数分别为SWI=0.924,SI=0.500,其中以开花习性最低(SWI=0.139,SI=0.014),而Shannon-Weaver指数以粒色最高为1.841,Simpson指数为0.712。利用48对SSR引物对这些材料进行了遗传多样性分析,结果如下:(1)在48对花生的SSR引物中,有35对(占所用引物总数的72.9%)具有多态性,共检测到215条多态性条带,平均每对引物可扩增6条多态性带。(2)根据SSR扩增结果对75份材料的聚类分析结果表明,这些材料的相似系数(GS)为0.25～0.85,平均GS值为0.55。28个审定品种的相似系数为0.39～0.85,平均为0.60。     

Genetic diversity among cultivars, landraces and wild relatives of rice as revealed by microsatellite markers

Genetic diversity among 35 rice accessions, which included 19 landraces, 9 cultivars and 7 wild relatives, was investigated by using microsatellite (SSR) markers distributed across the rice genome. The mean number of alleles per locus was 4.86, showing 95.2% polymorphism and an average polymorphism information content of 0.707. Cluster analysis based on microsatellite allelic diversity clearly demarcated the landraces, cultivars and wild relatives into different groups. The allelic richness computed for the clusters indicated that genetic diversity was the highest among wild relatives (0.436), followed by landraces (0.356), and the lowest for cultivars. Allelic variability among the SSR markers was high enough to categorize cultivars, landraces and wild relatives of the rice germplasm, and to catalogue the genetic variability observed for future use. The results also suggested the necessity to introgress genes from landraces and wild relatives into cultivars, for cultivar improvement.     

Genetic Diversity and Population Structure Among Oat Cultivars and Landraces

In this study, genetic diversity among 177 oat (Avena sativa L.) accessions including both white and red oat landraces and 36 commercial cultivars was studied for simple sequence repeat (SSR) loci. Thirty-one genomic and expressed sequence tags (EST)-derived primer pairs were selected according to high polymorphism from an initial 66 SSR batch. Markers revealed a high level of polymorphism, detecting a total of 454 alleles. The average gene diversity for the whole sample was 0.29. Genetic similarity, calculated using the Dice coefficient, was used for cluster analysis, and principal component analysis was also applied. In addition, population structure using a Bayesian clustering approach identified discrete subpopulation based on allele frequency and showed similar clustering of oat genotypes in four groups. Accessions could be classified into four main clusters that clearly separated the commercial cultivars, the red oat landraces and two clusters of white oat landraces. Cultivars showed less diversity than the landraces indicating a reduction of genetic diversity during breeding, whereas white oat landraces showed higher diversity than red ones. The average polymorphic information content of 0.80 for the SSR loci indicated the usefulness of many of the SSR for genotype identification. In particular, two markers, MAMA5 and AM04, with a total of 50 alleles and a high discrimination power (>0.90), were sufficient to discriminate among all commercial cultivars studied highlighting their potential use for variety identification.     

Genetic relationship among wild,landraces and cultivars of hazelnut (Corylus avellana) from Portugal revealed through ISSR and AFLP markers

The genus Corylus, a member of the birch family Betulaceae, includes several species that are widely distributed throughout temperate regions of the Northern Hemisphere. This study assesses the genetic diversity in 26 international cultivars and 32 accessions of Corylus avellana L. from Portugal: 13 wild genotypes and 19 landraces. The genetic relationships among the 58 hazelnuts (Corylus avellana L.) were analyzed using inter simple sequence repeat (ISSR) and amplified fragment length polymorphism (AFLP) markers. Eighteen ISSR primers and seven AFLP primer pairs generated a total of 570 unambiguous and repeatable bands, respectively, from which 541 (95.03 %) were polymorphic for both markers. Genetic similarity index values ranged from 0.239 for wild types and cultivars to 0.143 for landraces and wild types. The genetic relationships were presented as a Neighbor-Joining method dendrogram and a two-dimensional principal coordinate analysis (PCoA) plot. The Neighbor-Joining dendrogram showed three main clusters, and the PCoA analysis has shown to be congruent with the hierarchical analysis. Bayesian analysis clustered all individuals into three groups showing a good separation among wild genotypes, landraces and cultivars. The genetic diversity found on wild genotypes and Portuguese landraces may provide relevant information for the diversity conservation and it will be useful in breeding programs and to identify local selections for preservation.     

Genetic diversity in cultivars and landraces of Oryza sativa subsp. indica as revealed by AFLP markers.

Genetic diversity among 49 Indian accessions of rice (Oryza sativa subsp. indica), including 29 landraces from Jeypore, 12 modern cultivars, and 8 traditional cultivars from Tamil Nadu, was investigated using AFLP markers. In total, nine primer combinations revealed 664 AFLPs, 408 of which were found to be polymorphic. The percentage of polymorphic AFLPs was approximately the same within the cultivars and landraces. Similar results were obtained when genetic diversity values were estimated using the Shannon-Weiner index of diversity. Genetic diversity was slightly higher in the modern cultivars than in the traditional cultivars from Tamil Nadu. Among the landraces from Jeypore, the lowland landraces showed the highest diversity. The present study showed that the process of breeding modern cultivars did not appear to cause significant genetic erosion in rice. Cluster analysis and the first component of principle component analysis (PCA) both showed a clear demarcation between the cultivars and landraces as separate groups, although the genetic distance between them was narrow. The modern cultivars were positioned between the landraces from Jeypore and the traditional cultivars from Tamil Nadu. The second component of PCA further separated medium and upland landraces from lowland landraces, with the lowland landraces found closest to the traditional and modern cultivars.     

Differentiation and phylogenetic relationship among different cultivars of asparagus bean (Vigna unguiculata ssp. sesquipedalis) assessed using ISSR markers

Asparagus bean, one of the three subspecies of Vigna unguiculata, has a long cultivation history in China. The genetic diversity was analyzed based on the 66 landraces and cultivars cultivated in China by using ISSR molecular markers, with which 192 amplification loci were obtained 32.3% of them being polymorphic. The genetic differentiation analysis revealed a very high genetic diversity in the Chinese landraces, which confirmed that China is the secondary origin centre of the asparagus bean. The commercial cultivars bred in China were genetically highly homogenous, suggesting that the breeding process has resulted in disappearance of some of the genetic variation. A cluster analysis at 0.17 of Nei genetic distance divided the 66 landraces and cultivars into 9 groups. Their clustering pattern basically matches the phylogeny of those cultivars, and also corresponds to their geographical origin and morphological traits.     

Genetic diversity of allozymes in turnip (Brassica rapa L. var. rapa) from the Nordic area

Genetic diversity and relationships based on isozymes were studied in 31 accessions of turnip (Brassica rapa L. var. rapa). The material included varieties, elite stocks, landraces and older turnip of slash-and-burn type from the Nordic area. A total of 9 isozyme loci and 26 alleles were studied. The isozyme systems were ACO, DIA, GPI, GOT, PGM, PGD and SKD. The level of heterozygosity was reduced in the landraces, but it was high for the variety group 'Ostersundom'. Turnip has a higher genetic variation than other crops within B. rapa and than in other species with the same breeding system. The genetic diversity showed that 18.7% of the genetic variation was within the accessions, and the total H tau value was 0.358. Gpi-I and Pgd-I showed the lowest variation compared with the other loci. The cluster analysis revealed five clusters, with one main cluster including 25 of the 31 accessions. The dendrogram indicated that the variety group 'Ostersundom' clustered together whereas the variety group 'Bortfelder' was associated with country of origin. The landraces were spread in different clusters. The 'slash-and-burn' type of turnip belonged to two groups.     

Genetic Analysis of Indian Lentil (Lens culinaris Medikus) Cultivars and Landraces Using RAPD and STMS Markers

Molecular analysis of 29 lentil (Lens culinaris) cultivars and landraces of Indian origin was carried out using twenty RAPD and ten cross-species STMS primers. A total of 97 markers (72 RAPD and 25 STMS) were amplified of which 42.3% were polymorphic. Genetic similarity among the cultivars and landraces was 89.7%. The observed results indicated low level of genetic diversity in the studied material. UPGMA cluster analysis for the combined data of RAPD and STMS revealed two broad clusters — Cluster I with three landraces and Cluster II containing all remaining landraces and cultivars except Precoz. Germplasm line Precoz was found to be the most distinct in individual as well as combined analyses. All cultivars and landraces except K-75 and L4076 could be discriminated from one another using combined data for the two techniques. Germplasm lines Precoz, L830 and cultivars L4147 and JL3 were quite distinct and could be potential germplasm resource.     

Assessment of Genetic Diversity of Sweet Potato in Puerto Rico

Sweet potato (Ipomoea batatas L.) is the seventh most important food crop due to its distinct advantages, such as adaptability to different environmental conditions and high nutritional value. Assessing the genetic diversity of this important crop is necessary due to the constant increase of demand for food and the need for conservation of agricultural and genetic resources. In Puerto Rico (PR), the genetic diversity of sweet potato has been poorly understood, although it has been part of the diet since Pre-Columbus time. Thus, 137 landraces from different localities around PR were collected and subjected to a genetic diversity analysis using 23 SSR-markers. In addition, 8 accessions from a collection grown in Gurabo, PR at the Agricultural Experimental Station (GAES), 10 US commercial cultivars and 12 Puerto Rican accessions from the USDA repository collection were included in this assessment. The results of the analysis of the 23 loci showed 255 alleles in the 167 samples. Observed heterozygosity was high across populations (0.71) while measurements of total heterozygosity revealed a large genetic diversity throughout the population and within populations. UPGMA clustering method revealed two main clusters. Cluster 1 contained 12 PR accessions from the USDA repository collection, while cluster 2 consisted of PR landraces, US commercial cultivars and the PR accessions from GAES. Population structure analysis grouped PR landraces in five groups including four US commercial cultivars. Our study shows the presence of a high level of genetic diversity of sweet potato across PR which can be related to the genetic makeup of sweet potato, human intervention and out-crossing nature of the plant. The history of domestication and dispersal of sweet potato in the Caribbean and the high levels of genetic diversity found through this study makes sweet potato an invaluable resource that needs to be protected and further studied.     

Population Structure and Genetic Diversity in Popular Rice Varieties of India as Evidenced from SSR Analysis

We report here on the phylogenetic analysis, population substructure, and identification of molecular tags of 25 popular rice varieties and four landraces from different ecological belts of India employing a set of 52 simple sequence repeat (SSR) markers. Genetic analysis using the SSR markers categorized the genotypes into two major clusters, distributed according to their pedigree. Population structure analysis suggested that the optimum number of subpopulations was three (K?=?3) in the popular varieties and landraces. At K?=?5 the allelic distribution was much more similar to the phylogenetic dendrogram. The molecular diversity and population structure analysis indicated that there is not much variation among the popular rice cultivars of India. The study has identified SSR markers producing unique alleles, which should aid in the precise identification, maintenance, and genetic purity analysis of rice varieties.     

Genetic diversity of cultivated flax (Linum usitatissimum L.) germplasm assessed by retrotransposon-based markers

Retrotransposon segments were characterized and inter-retrotransposon amplified polymorphism (IRAP) markers developed for cultivated flax (Linum usitatissimum L.) and the Linum genus. Over 75 distinct long terminal repeat retrotransposon segments were cloned, the first set for Linum, and specific primers designed for them. IRAP was then used to evaluate genetic diversity among 708 accessions of cultivated flax comprising 143 landraces, 387 varieties, and 178 breeding lines. These included both traditional and modern, oil (86), fiber (351), and combined-use (271) accessions, originating from 36 countries, and 10 wild Linum species. The set of 10 most polymorphic primers yielded 141 reproducible informative data points per accession, with 52% polymorphism and a 0.34 Shannon diversity index. The maximal genetic diversity was detected among wild Linum species (100% IRAP polymorphism and 0.57 Jaccard similarity), while diversity within cultivated germplasm decreased from landraces (58%, 0.63) to breeding lines (48%, 0.85) and cultivars (50%, 0.81). Application of Bayesian methods for clustering resulted in the robust identification of 20 clusters of accessions, which were unstratified according to origin or user type. This indicates an overlap in genetic diversity despite disruptive selection for fiber versus oil types. Nevertheless, eight clusters contained high proportions (70?C100%) of commercial cultivars, whereas two clusters were rich (60%) in landraces. These findings provide a basis for better flax germplasm management, core collection establishment, and exploration of diversity in breeding, as well as for exploration of the role of retrotransposons in flax genome dynamics.