Origins,timing and introgression of domestic geese revealed by whole genome data

Background Geese are among the most important poultry species in the world.The current generally accepted hypothesis is that the European domestic geese originated from greylag geese(Anser anser),and Chinese domestic geese have two origins,most of which originated from swan geese(Anser cygnoides),and the Yili goose originated from greylag geese.To explain the origin and demographic history of geese,we selected 14 goose breeds from Europe and China and wild populations of swan and greylag geese,and whole genome sequencing data were obtained for 74 samples.Results Population structure analysis and phylogenetic trees showed that the wild ancestor of Chinese domestic geese,except for Yili,is the swan geese,and the wild ancestor of Chinese Yili and European domestic geese is greylag geese.Analysis of the demographic history suggests that the domestication of Chinese geese occurred~3499 years ago and that of the European geese occurred~7552 years ago.Furthermore,gene flow was observed between domestic geese and their wild ancestors.Analysis of introgression showed that Yili geese had been introgressed by Chinese domestic geese,and the body size of Yili geese may be influenced by introgression events of some growthrelated genes,including IGF-1.Conclusions Our study provides evidence for the origin of geese at the genome-wide level and advances the understanding of the history of goose domestication and the traits affected by introgression events.

Genome-wide recombination variation in biparental segregating and reciprocal backcross populations provides information for introgression breeding in Brassica napus

Variation in patterns of recombination in plant genomes provides information about species evolution,genetic diversity and crop improvement. We investigated meiotic crossovers generated in biparental segregating and reciprocal backcross populations of the allopolyploid genome of rapeseed(Brassica napus)(AACC, 2n = 38). A structured set of 1445 intercrossed lines was derived from two homozygous de novo genome-assembled parents that represented the major genetic clusters of semi-winter Chinese and winter European rapeseeds, and was used to increase QTL resolution and achieve genomic reciprocal introgression. A high-density genetic map constructed with 6161 genetic bins and anchored centromere regions was used to establish the pattern of recombination variation in each chromosome. Around 93%of the genome contained crossovers at a mean rate of 3.8 c M Mb^(-1), with the remaining 7% attributed to centromeres or low marker density. Recombination hotspots predominated in the A genome, including two-thirds of those associated with breeding introgression from B. rapa. Genetic background might affect recombination variation. Introgression of genetic diversity from European winter to Chinese semi-winter rapeseed showed an increase in crossover rate under the semi-winter environment. Evidence for an elevated recombination rate having historically contributed to selective trait improvement includes accumulation of favorable alleles for seed oil content on hotspots of chromosome A10. Conversely, strong artificial selection may affect recombination rate variation, as appears to be the case with a coldspot resulting from strong selection for glucosinolate alleles on A09. But the cold region would be promptly reactivated by crossing design indicated by the pedigree analysis. Knowledge of recombination hotspots and coldspots associated with QTL for 22 traits can guide selection strategies for introgression breeding between the two gene pools. These results and rich genomic resources broaden our understanding of recombination behavior in allopolyploids and may advance rapeseed genetic improvement.

Breeding by selective introgression: Theory, practices, and lessons learned from rice

Future demands for increased productivity and resilience to abiotic/biotic stresses of major crops require new technologies of breeding by design(BBD)built on massive information from functional and population genomics research.A novel strategy of breeding by selective introgression(BBSI)has been proposed and practiced for simultaneous improvement,genetic dissection and allele mining of complex traits to realize BBD.BBSI has three phases:a)developing large numbers of trait-specific introgression lines(ILs)using backcross breeding in elite genetic backgrounds as the material platform of BBD;b)efficiently identifying genes or quantitative trait loci(QTL)and mining desirable alleles affecting different target traits from diverse donors as the information platform of BBD;and c)developing superior cultivars by BBD using designed QTL pyramiding or marker-assisted recurrent selection.Phase(a)has been implemented massively in rice by many Chinese research institutions and IRRI,resulting in the development of many new green super rice cultivars plus large numbers of ILs in 30+elite genetic backgrounds.Phase(b)has been demonstrated in a series of proof-of-concept studies of high-efficiency genetic dissection of rice yield and tolerance to abiotic stresses using ILs and DNA markers.Phase(c)has also been implemented by designed QTL pyramiding,resulting in a prototype of BBD in several successful cases.The BBSI strategy can be easily extended for simultaneous trait improvement,efficient gene and QTL discovery and allele mining of complex traits using advanced breeding lines from crosses between a common"backbone"parent and a set of elite parents in conventional pedigree breeding programs.BBSI can be relatively easily adopted by breeding programs with small budgets,but the BBSI-based BBD strategy can be fully and more efficiently implemented by large seed companies with sufficient capacity.

Identification for Heat Tolerance in Backcross Recombinant Lines and Screening of Backcross Introgression Lines with Heat Tolerance at Milky Stage in Rice

The present study aimed at breeding new rice germplasms with similar genome but significantly differed in heat tolerance during the grain filling stage.A total of 791 BC1F8 backcross recombinant lines,derived from the cross of Xieqingzao B /N22 //Xieqingzao B,were used as materials.Each rice line was separated evenly into two groups,and the heat tolerance of all rice lines were evaluated at natural high temperature in fields.The rice lines with heat tolerant index higher than 90% or lower than 40% were selected to compare the phenotypic characters and further identify heat tolerance at the early milky stage in a phytotron.Rice lines with similar phenotypic characters but significantly differed in heat tolerance at the milky stage were analyzed by 887 simple sequence repeat markers that were evenly distributed on the 12 rice chromosomes.In the result,12(6 pairs) rice lines with similar phenotypic characters but significantly differed in heat tolerance at the milky stage were obtained.Molecular marker analysis indicated that the genomic polymorphism between 703T and 704S was the smallest in the 6 pairs of rice lines,with only 16 polymorphic sites,including 22 different alleles.The application of these two backcross introgression rice lines for future study on the mechanisms of heat tolerance in rice at the milky stage will be theoretically beneficial in reducing the interference caused by genetic differences from experimental materials.

Development and Identification of Introgression Lines from Cross of Oryza sativa and Oryza minuta

Introgression line population is effectively used in mapping quantitative trait loci(QTLs),identifying favorable genes,discovering hidden genetic variation,evaluating the action or interaction of QTLs in multiple conditions and providing the favorable experimental materials for plant breeding and genetic research.In this study,an advanced backcross and consecutive selfing strategy was used to develop introgression lines(ILs),which derived from an accession of Oryza minuta(accession No.101133) with BBCC genome,as the donor,and an elite indica cultivar IR24(O.sativa),as the recipient.Introgression segments from O.minuta were screened using 164 polymorphic simple sequence repeat(SSR) markers in the genome of each IL.Introgressed segments carried by 131 ILs covered the whole O.sativa genome.The average number of homozygous O.minuta segments per introgression line was about 9.99.The average length of introgressed segments was approximate 14.78 cM,and about 79.64% of these segments had sizes less than 20 cM.In the genome of each introgression line,the O.minuta chromosomal segments harbored chromosomal fragments of O.sativa ranging from 1.15% to 27.6%,with an overall average of 8.57%.At each locus,the ratio of substitution of O.minuta alleles had a range of 1.5% 25.2%,with an average of 8.3%.Based on the evaluation of the phenotype of these ILs,a wide range of alterations in morphological and yield-related traits were found.After inoculation,ILs 41,11 and 7 showed high resistance to bacterial blight,brown planthopper and whitebacked planthopper,respectively.These O.minuta-O.sativa ILs will serve as genetic materials for identifying and using favorable genes from O.minuta.

Dissection of heterosis for yield and related traits using populations derived from introgression lines in rice

Despite the great success achieved by the exploitation of heterosis in rice,the genetic basis of heterosis is still not well understood.We adopted an advanced-backcross breeding strategy to dissect the genetic basis of heterosis for yield and eight related traits.Four testcross(TC) populations with 228 testcross F1 combinations were developed by crossing57 introgression lines with four types of widely used male sterile lines using a North Carolina II mating design.Analysis of variance indicated that the effects of testcross F1 combinations and their parents were significant or highly significant for most of the traits in both years,and all interaction effects with year were significant for most of the traits.Positive midparent heterosis(HMP) was observed for most traits in the four TC populations in the two years.The relative HMPlevels for most traits varied from highly negative to highly positive.Sixty-two dominant-effect QTL were identified for HMPof the nine traits in the four TC populations in the two years.Of these,22 QTL were also identified for the performance of testcross F1.Most dominant-effect QTL could individually explain more than 10% of the phenotypic variation.Four QTL clusters were observed including the region surrounding the RM9–RM297 region on chromosome 1,the RM110–RM279–RM8–RM5699–RM452 region on chromosome 2,the RM5463 locus on chromosome 6 and the RM1146–RM147 region on chromosome 10.The identified QTL for heterosis provide valuable information for dissecting the genetic basis of heterosis.

Genome-wide prediction for hybrids between parents with distinguished difference on exotic introgressions in Brassica napus

Extensive exotic introgression could significantly enlarge the genetic distance of hybrid parental populations to promote strong heterosis.The goal of this study was to investigate whether genome-wide prediction can support pre-breeding in populations with exotic introgressions.We evaluated seed yield,seed yield related traits and seed quality traits of 363 hybrids of Brassica napus (AACC) derived from two parental populations divergent on massive exotic introgression of related species in three environments.The hybrids presented strong heterosis on seed yield,which was much higher than other investigated traits.Five genomic best linear unbiased prediction models considering the exotic introgression and different marker effects (additive,dominance,and epistatic effects) were constructed to test the prediction ability for different traits of the hybrids.The analysis showed that the trait complexity,exotic introgression,genetic relationship between the training set and testing set,training set size,and environments affected the prediction ability.The models with best prediction ability for different traits varied.However,relatively high prediction ability (e.g.,0.728 for seed yield) was also observed when the simplest models were used,excluding the effects of the special exotic introgression and epistasis effect by5-fold cross validation,which would simplify the prediction for the trait with complex architecture for hybrids with exotic introgression.The results provide novel insights and strategies for genome-wide prediction of hybrids between genetically distinct parent groups with exotic introgressions.

Introgressions from Oryza rufipogon Increase Photosynthetic Efficiency of KMR3 Rice Lines

To understand the wild Oryza genome effect on photosynthesis and its relation to total dry matter accumulation in an elite rice variety, a set of 40 stable introgression lines(ILs) BC_3F_8 derived from a cross of Oryza sativa(KMR3) × Oryza rufipogon(WR120) were grown under well watered conditions. Leaf gas exchange measurements and leaf chlorophyll estimates were conducted at the flowering stage. The results revealed significant variations in net photosynthetic rate(Pn), transpiration rate(E), transpiration efficiency(Pn/E) and carboxylation efficiency(Pn/C_i). Pn showed significant positive correlation with E, stomatal conductance(g_s), Pn/C_i and total canopy dry matter. Specific leaf area and leaf thickness were not significantly correlated with Pn. Thirty-seven out of 40 ILs showed higher Pn than KMR3 [11.28 μmol/(m^2·s)], and 20 ILs showed higher Pn than WR120 [15.08 μmol/(m^2·s)]. The line IL194 showed the highest Pn [21.62 μmol/(m^2·s)] with increased total canopy dry matter followed by lines IL381, IL106, IL363-12, IL198, IL86-18 and IL50, which exhibited Pn above 18.0 μmol/(m^2·s). The ILs with enhanced Pn are a potential source for developing rice varieties and hybrids with higher biomass and yield.

QTL underlying iron and zinc toxicity tolerances at seedling stage revealed by two sets of reciprocal introgression populations of rice(Oryza sativa L.)

Iron and zinc are two trace elements that are essential for rice. But they are toxic at higher concentrations, leading to severe rice yield losses especially in acid soils and inland valleys. In this study, two reciprocal introgression line(IL) populations sharing the same parents were used with high-density SNP bin markers to identify QTL tolerant to iron and zinc toxicities. The results indicated that the japonica variety 02,428 had stronger tolerance to iron and zinc toxicities than the indica variety Minghui 63. Nine and ten QTL contributing to iron and zinc toxicity tolerances,respectively, were identified in the two IL populations. The favorable alleles of most QTL came from 02,428. Among them, q FRRDW2, q ZRRDW3, and q FRSDW11 appeared to be independent of genetic background. The region C11S49–C11S60 on chromosome 11 harbored QTL affecting multiple iron and zinc toxicity tolerance-related traits, indicating partial genetic overlap between the two toxicity tolerances. Our results provide essential information and materials for developing excellent rice cultivars with iron and/or zinc tolerance by marker-assisted selection(MAS).

Introgression of Gossypium klotzschianum Genome into Cultivated Cotton,G.hirsutum

Exotic Gossypium germplasm is a rich source of useful agronomic traits for improvement programs of cultivated cotton.Efficient use of genetic variation available in the wild relatives depends

Yield Traits and Associated Marker Segregation in Elite Introgression Lines Derived from O. sativa × O. nivara

Introgression lines(ILs)derived from interspecific crosses are a source of new genetic variability.A total of 55 ILs derived from two crosses Swarna×O.nivara IRGC81848(population A)and Swarna×O.nivara IRGC81832(population B)were characterized for yield and yield-related traits/QTLs.Segregation of 103 simple sequence repeat(SSR)markers associated with yield-related QTLs was studied.Population A showed an average of 12.6%homozygous O.nivara alleles and population B showed 10.6%.Interestingly,three SSR markers,RM223,RM128 and RM517,showed conspicuous pattern of segregation.The distribution of parental alleles at three loci RM223,RM128 and RM517 linked to yield-related traits was unique.These markers flanked to several yield-related QTLs.RM223,flanking to qyld8.3,was heterozygous in almost all the 55 ILs except in IL10-3S and IL131S.RM128 on chromosome 1 and RM517 on chromosome 3 were mutually exclusive in 46 out of 55 ILs.These 46 ILs showed either of the marker allele RM128 or RM517 from O.nivara but not both.IL166S had both RM128and RM517 from O.nivara and the other ILs showed homozygous Swarna allele at RM517 except IL65S.Population structure assigned the 55 ILs to three sub-populations based on their genomic diversity.IL65S,IL166S,IL248S,IL7K and IL250K showed high yields in multi-location trials,and IL248S was released for cultivation as DRRDhan 40.

Characterization of Newly Developed Wheat/Barley Introgression Lines in Respect of Aluminium Tolerance

The Al tolerance of newly developed wheat/barley disomic addition, substitution and translocation lines carry chromosomes of three different barley cultivars was evaluated by comparing the root growth in solution containing 75 μM AlCl3 at pH 4.0 to that of known Al-tolerant and sensitive wheat genotypes. The wheat Asakaze komugi, barley Manas cultivars and their hybrid derivatives were found to have high levels of Al tolerance. The wheat line Mv9kr1, barley cultivar Igri and progenies of the hybrids were sensitive to Al. In most cases, the Al tolerance of the wheat/barley introgression lines derived from Al-sensitive wheat Mv9kr1 and barley Betzes with moderate Al tolerance was similar to that of the wheat parents, but the 2DS.2DL-1HS translocation line of Mv9kr1/Betzes exhibited more intensive root growth, while accumulating less Al than the parental lines. This indicates that either the lack of the distal part of chromosome 2DL or the presence of the distal part of 1HS improved the Al tolerance level.

Hominin Evolution Was Caused by Introgression from Gorilla

The discovery of Paranthropus deyiremeda in 3.3 - 3.5 million-year-old fossil sites in Afar, together with 30% of the gorilla genome showing lineage sorting between humans and chimpanzees, and a NUMT (“nuclear mitochondrial DNA segment”) on chromosome 5 that is shared by both gorillas, humans and chimpanzees, and shown to have diverged at the time of the Pan-Homo split rather than the Gorilla/Pan-Homo split, provides conclusive evidence that introgression from the gorilla lineage caused the Pan-Homo split, and the speciation of both the Australopithecus lineage and the Paranthropus lineage.

Preliminary Phenotypic and SNP-Based Molecular Characterization of Maize (<i>Zea mays</i>L.)-Mexicana (<i>Zea mays</i>SSP. <i>Mexicana</i>) Introgression Lines under Inbred Background of 48-2

Wild relatives possess potential genetic diversity for maize (Zea mays L.) improvement. Characterization of maize-mexicana introgression lines (ILs) is of great value to diversify the genetic base and improve the maize germplasm. Four maize-mexicana IL generations, i.e. BC1, BC2, BC3, and RIL, were constructed under the elite inbred background of 48-2, elite inbred line that is widely used in maize breeding in Southwestern China, and were phenotyped in different years and genotyped with 56110 SNPs. The results indicated that 48-2 had higher phenotypic performances than all the characterized ILs on most of the agronomic traits. Compared with other ILs, BC2 individuals exhibited more similar performance to 48-2 on most traits and possessed the highest kernel ratio (66.5%). Population structure and principal component analysis indicated that BC3 individuals gathered closer to 48-2 and exhibited the lowest mexicana-introgression frequency (0.50%), while BC2 (29.06%) and RIL (18.52%) showed higher introgression frequency. The high level of genetic diversity observed in the maize-mexicana ILs demonstrated that Z. mays ssp. mexicana can serve as a potential source for the enrichment of maize germplasm.

Introgression of Bt Genes in Novel Germplasm and Contribution to Indian Cotton Economy

Emergence of transgenic Bt-cotton technology has opened up a new chapter in Indian cotton production in 21st century.The cry1Ac gene of Monsanto derived from American Upland Coker-312 background was not directly suitable for varied cotton growing situations in India.Delivery of

Introgression of Two Drought QTLs into FUNAABOR-2 Early Generation Backcross Progenies Under Drought Stress at Reproductive Stage

FUNAABOR-2 is a popular Ofada rice variety grown in a large area under rainfed upland condition across western states of Nigeria. We used the combination of phenotypic and marker-assisted selection(MAS) to improve grain yield of FUNAABOR-2 under drought stress(DS) at the reproductive stage via introgression of two drought quantitative trait loci(QTLs), qDTY12.1 and qDTY2.3. Foreground selection was carried out using peak markers RM511 and RM250, associated with qDTY12.1 and qDTY2.3, respectively, followed by recombinant selection with RM28099 and RM1261 distally flanking qDTY12.1. Furthermore, BC1 F2-derived introgressed lines and their parents were evaluated under DS and non-stress(NS) conditions during the 2015–2016 dry season. Overall reduction of grain yield under DS compared to NS was recorded. Introgressed lines with qDTY12.1 and qDTY2.3 combinations showed higher yield potential compared to lines with single or no QTL under DS, indicating significant positive interactions between the two QTLs under the FUNAABOR-2 genetic background. Pyramiding of qDTY12.1 and qDTY2.3 in the FUNAABOR-2 genetic background led to higher grain yield production under DS and NS.

Utilizing resequencing big data to facilitate Brassica vegetable breeding:tracing introgression pedigree and developing highly specific markers for clubroot resistance

Clubroot caused by Plasmodiophora brassicae is a devastating disease of Cruciferous crops.Developing cultivars with clubroot resistance(CR)is the most effective control measure.For the two major Brassica vegetable species B.rapa and B.oleracea,several commercial cultivars with unclear CR pedigrees have been intensively used as CR donors in breeding.However,the continuous occurrence of CR-breaking makes the CR pedigree underlying these cultivars one of the breeders'most urgent concerns.The complex intraspecific diversity of these two major Brassica vegetables has also limited the applicability of CR markers in different breeding programs.Here we first traced the pedigree underlying two kinds of CR that have been widely applied in breeding by linkage and introgression analyses based on public resequencing data.In B.rapa,a major locus CRzi8 underlying the CR of the commercial CR donor‘DegaoCR117’was identified.CRzi8 was further shown to have been introgressed from turnip(B.rapa ssp.rapifera)and that it carried a potential functional allele of Crr1a.The turnip introgression carried CRb^(c),sharing the same coding sequence with the CRb that was also identified from chromosome C07 of B.oleracea CR cultivars with different morphotypes.Within natural populations,variation analysis of linkage intervals of CRzi8,PbBa8.1,CRb,and CRb^(c)yielded easily resolved InDel markers(>20 bp)for these fundamental CR genes.The specificity of these markers was tested in diverse cultivars panels,and each exhibited high reliability in breeding.Our research demonstrates the value of the practice of applying resequencing big data to solve urgent concerns in breeding programs.

Climate-influenced boreotropical survival and rampant introgressions explain the thriving of New World grapes in the north temperate zone

The north temperate region was characterized by a warm climate and a rich thermophilic flora before the Eocene,but early diversifications of the temperate biome under global climate change and biome shift remain uncertain.Moreover,it is becoming clear that hybridization/introgression is an important driving force of speciation in plant diversity.Here,we applied analyses from biogeography and phylogenetic networks to account for both introgression and incomplete lineage sorting based on genomic data from the New World Vitis,a charismatic component of the temperate North American flora with known and suspected gene flow among species.Biogeographic inference and fossil evidence suggest that the grapes were widely distributed from North America to Europe during the Paleocene to the Eocene,followed by widespread extinction and survival of relicts in the tropical New World.During the climate warming in the early Miocene,a Vitis ancestor migrated northward from the refugia with subsequent diversification in the North American region.We found strong evidence for widespread incongruence and reticulate evolution among nuclear genes within both recent and ancient lineages of the New World Vitis.Furthermore,the organellar genomes showed strong conflicts with the inferred species tree from the nuclear genomes.Our phylogenomic analyses provided an important assessment of the wide occurrence of reticulate introgression in the New World Vitis,which potentially represents one of the most important mechanisms for the diversification of Vitis species in temperate North America and even the entire temperate Northern Hemisphere.The scenario we report here may be a common model of temperate diversification of flowering plants adapted to the global climate cooling and fluctuation in the Neogene.

Deep genome skimming reveals the hybrid origin of Pseudosasa gracilis (Poaceae: Bambusoideae)

Pseudosasa gracilis(Poaceae:Bambusoideae)is a temperate woody bamboo species endemic to Southcentral China with a narrow distribution.Previous phylogenetic studies revealed an unexpected,isolated phylogenetic position of Ps.gracilis.Here we conducted phylogenomic analysis by sampling populations of Ps.gracilis and its sympatric species Ps.nanunica and Sinosasa polytricha reflecting different genomic signals,by deep genome skimming.Integrating molecular evidence from chloroplast genes and genome-wide SNPs,we deciphered the phylogenetic relationships of Ps.gracilis.Both plastid and nuclear data indicate that Ps.gracilis is more closely related to Sinosasa,which is discordant with the taxonomic treatment.To further explore this molecular-morphological conflict,we screened 411“perfect-copy”syntenic genes to reconstruct phylogenies using both the concatenation and coalescent methods.We observed extensive discordance between gene trees and the putative species tree.A significant hybridization event was detected based on 411 genes from the D subgenome,showing Ps.gracilis was a hybrid descendant between Sinosasa longiligulata and Ps.nanunica,with 63.56%and 36.44%inheritance probabilities of each parent.Moreover,introgression events were detected in the C subgenome between Ps.gracilis and S.polytricha in the same distribution region.Our findings suggest that sympatric hybridization and introgression play a crucial role in the origin of Ps.gracilis.By providing an empirical example of bamboo of hybrid origin using comprehensive analyses based on genomic data from different inheritance systems and morphological characters,our study represents a step forward in understanding of reticulate evolution of bamboos.

Genomic atlases of introgression and differentiation reveal breeding footprints in Chinese cultivated rice

The long history of cultivation and breeding has left a variety of footprints in the genomes of Asian cultivated rice(Oryza sativa L.).In this study,we focus on two types of genomic footprints,introgression and differentiation,in a population of more than 1200 Chinese rice accessions.We found that a Xian/indica and a temperate Geng/japonica accession respectively contained an average of 19.3-Mb and 6.8-Mb alien introgressed chromosomal segments,of which many contained functional sequence variants,quantitative trait loci,or genes controlling flowering,grain,and resistance traits.Notably,we found most introgressions,including the known heterotic loci Hd3 a and TAC1,were distributed differentially between the female and male parents of three-line indica hybrid rice,indicating their potential contribution to heterosis.We also found many differentiated regions between subgroups within a subpopulation contained agronomically important loci,such as DTH7,Hd1 for heading date,and qCT7 for cold tolerance,providing new candidates for studying local adaptation or heterosis.Tracing these footprints allows us to better understand the genetic exchange or differentiation underlying agronomic traits in modern Chinese rice cultivars.These findings also provide potential targets for rice genetic research and breeding.