Soybean yield has traditionally been increased through high planting density,but investigating plant height and petiole traits to select for compact architecture,lodging resistance,and high yield varieties is an under...Soybean yield has traditionally been increased through high planting density,but investigating plant height and petiole traits to select for compact architecture,lodging resistance,and high yield varieties is an underexplored option for further improving yield.We compared the relationships between yield-related traits,lodging resistance,and petioleassociated phenotypes in the short petiole germplasm M657 with three control accessions during 2017–2018 in four locations in the Huang–Huai region,China.The results showed that M657 exhibited stable and high tolerance to high planting density and resistance to lodging,especially at the highest density(8×105 plants ha–1).The regression analysis indicated that a shorter petiole length was significantly associated with increased lodging resistance.The yield analysis showed that M657 achieved higher yields under higher densities,especially in the northern part of the Huang–Huai region.Among the varieties,there were markedly different responses to intra-and inter-row spacing designs with respect to both lodging and yield that were related to location and density.Lodging was positively correlated with planting density,plant height,petiole length,and number of effective branches,but negatively correlated with stem diameter,seed number per plant,and seed weight per plant.The yield of soybean was increased by appropriately increasing the planting density on the basis of the current soybean varieties in the Huang–Huai region.This study provides a valuable new germplasm resource for the introgression of compact architecture traits that are amenable to providing a high yield in high density planting systems,and it establishes a high-yield model of soybean in the Huang–Huai region.展开更多
The objective of this study was to investigate the correlations between antioxidations and the contents of the total phenolics and anthocyanin in 127 accessions of black soybean. A T-test, a fast clustering procedure,...The objective of this study was to investigate the correlations between antioxidations and the contents of the total phenolics and anthocyanin in 127 accessions of black soybean. A T-test, a fast clustering procedure, and a correlation coefficient analysis were used for experimentation. The variation ranges of the total antioxidant capacity （TAC）, the total phenolics, and anthocyanin contents in 127 black soybean accessions were 0.44-3.56, 7.05-74.82, and 0.22-1.87 mg g-l, respectively, displaying significant genotype differences. The major differences in TAC, the total phenolics, and the anthocyanin contents existed among various types of accessions from geographical regions. The differences between the accessions from black and yellow soybeans, spring and autumn, summer and autumn, Dongbeichun and Nanfangchun, Dongbeichun and Nanfangxia, Beifangchun and Nanfangchun, and Beifangchun and Nanfangxia were significant at 0.01 or 0.05 levels, respectively. The general tendency was that the TAC, the total phenolics, and the anthocyanin contents of Beifangchun accessions were higher than that of Dongbeichun ones, while that of Nanfangchun accessions were the worst. 127 black soybean accessions could be clustered into 6 clusters, which consisted of 3, 24, 20, 31, 37, and 12 accessions, respectively. The most significant （P 〈 0.01） correlations existed respectively between the TAC and the total showed that the total substances phenolics content, and the TAC and the anthocyanin content of black soybean. The results phenolics and anthocyanin in black soybean seed coat were the important antioxidation展开更多
Rhizosphere colonization is a key requirement for the application of plant growth-promoting rhizobacteria(PGPR)as a bioferilizer.Signaling molecules are often exchanged between PGPR and plants,and genes in plants may ...Rhizosphere colonization is a key requirement for the application of plant growth-promoting rhizobacteria(PGPR)as a bioferilizer.Signaling molecules are often exchanged between PGPR and plants,and genes in plants may respond to the action of PGPR.Here,the luciferase luxAB gene was electrotransformed into Pseudomonas sp.strain TK35,a PGPR with an afinity for tobacco,and the labelled TK35(TK35-L)was used to monitor colonization dynamics in the tobacco rhizosphere and evaluate the effects of colonization on tobacco growth and root development.The transcript levels of the hydroxyproline rich glycoprotein HRGPnt3 gene,a lateral root induction indicator,in tobacco roots were examined by qPCR.The results showed that TK35-L could survive for long periods in the tobacco rhizosphere and colonize new spaces in the tobacco rhizosphere following tobacco root extension,exhibiting significant increases in root development,seedling growth and potassium accumulation in tobacco plants.The upregulation of HRGPnt3 transcription in the inoculated tobacco suggested that TK35-L can promote tobacco root development by upregulating the transcript levels of the HRGPnt3 gene,which promotes tobacco seedling growth.These findings lay a foundation for future studies on the molecular mechanism underlying the plant growth-promoting activities of PGPR.Futhermore,this work provided an ideal potential strain for biofertilizer production.展开更多
Soybean is a widely planted genetically modified crop around the world. However, it is still one of the most recalcitrant crops for genetic transformation due to the difficulty of regeneration via organogenesis and so...Soybean is a widely planted genetically modified crop around the world. However, it is still one of the most recalcitrant crops for genetic transformation due to the difficulty of regeneration via organogenesis and some factors that affect the transformation efficiency. The percentages of resistant bud formation and transient expression efficiency are important indexes reflecting the regeneration and transformation efficiency of soybean. In this study, the percentages of resistant bud formation and transient expression of 13-glucuronidase （GUS） were compared after treatment with sonication or surfactant and co-treatment with both. The results showed that treatment with either sonication or surfactant increased the percentage of resistant bud formation and transient expression efficiency. The highest percentages were acquired and significantly improved when cotyledon node explants were co-treated with sonication for 2 s and surfactant at 0.02% （v：v） using two different soybean genotypes, Jack and Zhonghuang 10. The improved transformation efficiency of this combination was also evaluated by development of herbicide-tolerant soybeans with transformation efficiency at 2.5-5.7% for different geno- types, which was significantly higher than traditional cotyledonary node method in this study. These results suggested that co-treatment with surfactant and sonication significantly improved the percentages of resistance bud formation, transient expression efficiency and stable transformation efficiency in soybean.展开更多
Developing expressed sequence tag-derived SSR （EST-SSR） markers is imperative in genetic research. In this paper, we reported 37 EST-SSR markers which were developed from 286 unigenes obtained from soybean cDNA libr...Developing expressed sequence tag-derived SSR （EST-SSR） markers is imperative in genetic research. In this paper, we reported 37 EST-SSR markers which were developed from 286 unigenes obtained from soybean cDNA library. Among the 286 markers designed for the 4 accessions of Glycine max and 6 of its wild progenitor （G. soja） within the subgenus Soja, 209 markers amplified DNA fragments, taking 73.1% and 37 markers appeared to be polymorphic, which was 12.9% of the total. The 37 loci detected a total of 142 alleles, while the PIC values varied from 0.194 to 0.794. Both the number of alleles per locus and PIC value were significantly related to the SSR motif. Six EST-SSR loci may be fixed for different alleles between G. max and G. soja since they were particularly polymorphic among the 6 G. soja accessions. A neighbor-joining tree placed the G. max accessions together as a group within the G. soja, though the average genetic distance among G. soja accessions was much higher. These new EST-SSRs markers will be useful for genetic diversity analysis, genetic mapping construction and gene discovery in Soja subgenus.展开更多
Information on the center of genetic diversity of soybean（Glycine max） will be helpful not only for designing efficient strategies for breeding programs, but also for understanding the domestication and origin of th...Information on the center of genetic diversity of soybean（Glycine max） will be helpful not only for designing efficient strategies for breeding programs, but also for understanding the domestication and origin of this species. Here, we describe an analysis of genetic diversity based on simple-sequence repeat（SSR） variations within a core collection of 2 111 accessions of Chinese soybean landraces. Prior to the diversity assessment, the geographic origin of each accession was mapped. The map was then divided into grids each 2.5° in latitude and 5° in longitude. We found two regions that had higher number of alleles（NA） and greater polymorphic information content（PIC） values than the others. These regions are adjacently located within grid position of 30°–35°N×105°–110°E, which includes the valley of the middle and lower reaches of the Wei River, and the valley of the upper reaches of the Hanjiang River. It was also observed that in many regions, genetic diversity decreased with the increase in distance from the center. Another region, in northern Hebei Province（115°–120°E×40°–42.5°N）, was observed having higher diversity than any surrounding regions, indicating that this is a sub-center of soybean diversity. Based on the presented results, the domestication and origin of soybean are also discussed.展开更多
The growth periods（GPs, from planting/emergence to reproductive stage 8（R8） of soybean cultivars vary in different ecological regions, especially in China with a very complex soybean cropping system. In this study,...The growth periods（GPs, from planting/emergence to reproductive stage 8（R8） of soybean cultivars vary in different ecological regions, especially in China with a very complex soybean cropping system. In this study, a 3-yr experimental study was undertaken in three geographical locations of China from 2008 to 2010, including the Northeast（40.66-45.85°N）, Huang-Huai（34.75-38.04°N） and southern（22.82-30.60°N） eco-regions with about 250 accessions in each region to clarify the classification of maturity group（MG） and identify the cultivars with stable GP to increase the knowledge about the GP distribution of soybean cultivars in China. GPs of soybean cultivars in different eco-regions were significant different with a gradual decrease from 115-125 d in the Northeast part to the 85-100 d in the southern part of China. The geographical location was the major factor for GP of cultivars from the Northeast, while the year of planting was the major factor affecting the stability of GPs in Huang-Huai summer and southern summer soybean. AMMI2（additive main effects and multiplicativeinteraction）-Biplot analysis showed that the GPs of soybean cultivars from the Northeast eco-region have a comparatively satisfactory environmental stability. Moreover, soybean cultivars with moderate GP/MG and stable environment adaptability in different eco-regions were identified based on the linear regression and AMMI analysis, which was important for the accurate classification of soybean MGs in future. Taken together, our results reflected the genetic diversity, geographical distribution and environmental stability of the Chinese soybean GP trait. Soybean cultivars with stable GP for various Chinese eco-regions would be beneficial for Chinese soybean genetic improvement, varietal introduction, exchange, and soybean breeding program for wide adaptability.展开更多
The leaf is the main photosynthetic organ of plants,and it plays a significant role in the yield of crop species.Identifying the causal mutations and candidate genes that underlie leaf phenotypic variation is an impor...The leaf is the main photosynthetic organ of plants,and it plays a significant role in the yield of crop species.Identifying the causal mutations and candidate genes that underlie leaf phenotypic variation is an important breeding target in soybean grain yield improvement.An ethyl methyl sulfonate(EMS)-induced soybean mutant DWARFCRINKLEDLEAF1(DCL1)with an aberrant crinkled leaf phenotype was identified in the background of the soybean cultivar Zhongpin 661(Zp661).We constructed an F2 segregating population from a cross between Zp661 and DCL1 in order to investigate the genomic locus associated with the crinkled leaf trait.Using bulk segregant analysis(BSA)combined with the whole-genome resequencing method,the Euclidean distance(ED)correlation algorithm detected 12 candidate genomic regions with a total length of 20.32 Mb that were linked to the target trait.Following a comparative analysis of the sequence data for the wild-type and mutant pools,only one single nucleotide mutation(C:G>T:A)located on the first exon of Glyma.19G207100 was found to be associated with the trait.Candidate gene validation based on a CAPS marker derived from the detected single-nucleotide polymorphism(SNP)indicated a nucleotide polymorphism between the two parents.Therefore,our findings reveal that Glyma.19G207100,which is renamed as GLYCINE MAX DWARF CRINKLED LEAF 1(GmDCL1),is a promising candidate gene involved in the morphogenesis of the crinkled leaf trait of the soybean mutant DCL1.This study provides a basis for the functional validation of this gene,with prospects for soybean breeding targeting grain yield enhancement.展开更多
For clarifying the hierarchical patterns of population structure of soybean landraces in China, the seven clusters previously identified using Bayesian clustering of 1 504 soybean landraces based on SSR markers genoty...For clarifying the hierarchical patterns of population structure of soybean landraces in China, the seven clusters previously identified using Bayesian clustering of 1 504 soybean landraces based on SSR markers genotyping data were further analyzed. Using the largest value of AK, these landraces could be split into 20 sub-clusters, which was supported by highly significant pairwise Fst-values and generally in accordance with the geographic origin and sowing types. The autumn-sowing types ended up in one distinct sub-cluster from the otherwise summer-sowing type, where the autumn- sowing types are most likely derived from. The division into 20 sub-clusters explained 7.3% of the genetic variation, next to 9.7% present among the seven clusters, 81.1% residing among landraces within sub-clusters, and 1.9% within the landraces. The distribution pattern of genetic diversity among the sub-clusters of each cluster was uneven, with two HSuM sub-clusters （Central China） and some South China sub-clusters showing significantly higher level of genetic diversity.展开更多
基金funded by the National Natural Science Foundation of China (31271753)the Central Publicinterest Scientific Institution Basal Research Fund, China (S2021ZD02)the Agricultural Science and Technology Innovation Program (ASTIP) of the Chinese Academy of Agricultural Sciences (CAAS-ZDRW202003-1)。
文摘Soybean yield has traditionally been increased through high planting density,but investigating plant height and petiole traits to select for compact architecture,lodging resistance,and high yield varieties is an underexplored option for further improving yield.We compared the relationships between yield-related traits,lodging resistance,and petioleassociated phenotypes in the short petiole germplasm M657 with three control accessions during 2017–2018 in four locations in the Huang–Huai region,China.The results showed that M657 exhibited stable and high tolerance to high planting density and resistance to lodging,especially at the highest density(8×105 plants ha–1).The regression analysis indicated that a shorter petiole length was significantly associated with increased lodging resistance.The yield analysis showed that M657 achieved higher yields under higher densities,especially in the northern part of the Huang–Huai region.Among the varieties,there were markedly different responses to intra-and inter-row spacing designs with respect to both lodging and yield that were related to location and density.Lodging was positively correlated with planting density,plant height,petiole length,and number of effective branches,but negatively correlated with stem diameter,seed number per plant,and seed weight per plant.The yield of soybean was increased by appropriately increasing the planting density on the basis of the current soybean varieties in the Huang–Huai region.This study provides a valuable new germplasm resource for the introgression of compact architecture traits that are amenable to providing a high yield in high density planting systems,and it establishes a high-yield model of soybean in the Huang–Huai region.
基金This work was supported by the National Natural Science Foundation of China （30200171）.
文摘The objective of this study was to investigate the correlations between antioxidations and the contents of the total phenolics and anthocyanin in 127 accessions of black soybean. A T-test, a fast clustering procedure, and a correlation coefficient analysis were used for experimentation. The variation ranges of the total antioxidant capacity （TAC）, the total phenolics, and anthocyanin contents in 127 black soybean accessions were 0.44-3.56, 7.05-74.82, and 0.22-1.87 mg g-l, respectively, displaying significant genotype differences. The major differences in TAC, the total phenolics, and the anthocyanin contents existed among various types of accessions from geographical regions. The differences between the accessions from black and yellow soybeans, spring and autumn, summer and autumn, Dongbeichun and Nanfangchun, Dongbeichun and Nanfangxia, Beifangchun and Nanfangchun, and Beifangchun and Nanfangxia were significant at 0.01 or 0.05 levels, respectively. The general tendency was that the TAC, the total phenolics, and the anthocyanin contents of Beifangchun accessions were higher than that of Dongbeichun ones, while that of Nanfangchun accessions were the worst. 127 black soybean accessions could be clustered into 6 clusters, which consisted of 3, 24, 20, 31, 37, and 12 accessions, respectively. The most significant （P 〈 0.01） correlations existed respectively between the TAC and the total showed that the total substances phenolics content, and the TAC and the anthocyanin content of black soybean. The results phenolics and anthocyanin in black soybean seed coat were the important antioxidation
基金Supported by the National Natural Science Foundation of China(41401269)the Key Project of the University Natural Science Research Project of Anhui Province,China(KJ2019A0183)+1 种基金the Open Fund of Anhui Province Key Laboratory of Farmland Ecological Conservation and Pollution Prevention(FECPP201902)the Key Research Project of China National Tobacco Corporation Hubei Company(027Y2020-011).
文摘Rhizosphere colonization is a key requirement for the application of plant growth-promoting rhizobacteria(PGPR)as a bioferilizer.Signaling molecules are often exchanged between PGPR and plants,and genes in plants may respond to the action of PGPR.Here,the luciferase luxAB gene was electrotransformed into Pseudomonas sp.strain TK35,a PGPR with an afinity for tobacco,and the labelled TK35(TK35-L)was used to monitor colonization dynamics in the tobacco rhizosphere and evaluate the effects of colonization on tobacco growth and root development.The transcript levels of the hydroxyproline rich glycoprotein HRGPnt3 gene,a lateral root induction indicator,in tobacco roots were examined by qPCR.The results showed that TK35-L could survive for long periods in the tobacco rhizosphere and colonize new spaces in the tobacco rhizosphere following tobacco root extension,exhibiting significant increases in root development,seedling growth and potassium accumulation in tobacco plants.The upregulation of HRGPnt3 transcription in the inoculated tobacco suggested that TK35-L can promote tobacco root development by upregulating the transcript levels of the HRGPnt3 gene,which promotes tobacco seedling growth.These findings lay a foundation for future studies on the molecular mechanism underlying the plant growth-promoting activities of PGPR.Futhermore,this work provided an ideal potential strain for biofertilizer production.
基金supported by the National Tran sgenic Major Program of China (2014ZX08004001)
文摘Soybean is a widely planted genetically modified crop around the world. However, it is still one of the most recalcitrant crops for genetic transformation due to the difficulty of regeneration via organogenesis and some factors that affect the transformation efficiency. The percentages of resistant bud formation and transient expression efficiency are important indexes reflecting the regeneration and transformation efficiency of soybean. In this study, the percentages of resistant bud formation and transient expression of 13-glucuronidase （GUS） were compared after treatment with sonication or surfactant and co-treatment with both. The results showed that treatment with either sonication or surfactant increased the percentage of resistant bud formation and transient expression efficiency. The highest percentages were acquired and significantly improved when cotyledon node explants were co-treated with sonication for 2 s and surfactant at 0.02% （v：v） using two different soybean genotypes, Jack and Zhonghuang 10. The improved transformation efficiency of this combination was also evaluated by development of herbicide-tolerant soybeans with transformation efficiency at 2.5-5.7% for different geno- types, which was significantly higher than traditional cotyledonary node method in this study. These results suggested that co-treatment with surfactant and sonication significantly improved the percentages of resistance bud formation, transient expression efficiency and stable transformation efficiency in soybean.
基金supported by the National High-Tech R&D Program of China(863Program,2006AA10A110 and 2006AA10Z164)National Basic Research Program of China(973 Program,2010CB125900 and 2004CB117203)the Academy and Institute Foundation for Basic Scientific Research in Institute of Crop Science,Chinese Academy of Agricultural Sciences
文摘Developing expressed sequence tag-derived SSR （EST-SSR） markers is imperative in genetic research. In this paper, we reported 37 EST-SSR markers which were developed from 286 unigenes obtained from soybean cDNA library. Among the 286 markers designed for the 4 accessions of Glycine max and 6 of its wild progenitor （G. soja） within the subgenus Soja, 209 markers amplified DNA fragments, taking 73.1% and 37 markers appeared to be polymorphic, which was 12.9% of the total. The 37 loci detected a total of 142 alleles, while the PIC values varied from 0.194 to 0.794. Both the number of alleles per locus and PIC value were significantly related to the SSR motif. Six EST-SSR loci may be fixed for different alleles between G. max and G. soja since they were particularly polymorphic among the 6 G. soja accessions. A neighbor-joining tree placed the G. max accessions together as a group within the G. soja, though the average genetic distance among G. soja accessions was much higher. These new EST-SSRs markers will be useful for genetic diversity analysis, genetic mapping construction and gene discovery in Soja subgenus.
基金supported by the National Basic Research Program of China(973,G1998010203 and 2004CB117203)the Agricultural Science and Technology Innovation Program(ASTIP)of Chinese Academy of Agricultural Sciences
文摘Information on the center of genetic diversity of soybean（Glycine max） will be helpful not only for designing efficient strategies for breeding programs, but also for understanding the domestication and origin of this species. Here, we describe an analysis of genetic diversity based on simple-sequence repeat（SSR） variations within a core collection of 2 111 accessions of Chinese soybean landraces. Prior to the diversity assessment, the geographic origin of each accession was mapped. The map was then divided into grids each 2.5° in latitude and 5° in longitude. We found two regions that had higher number of alleles（NA） and greater polymorphic information content（PIC） values than the others. These regions are adjacently located within grid position of 30°–35°N×105°–110°E, which includes the valley of the middle and lower reaches of the Wei River, and the valley of the upper reaches of the Hanjiang River. It was also observed that in many regions, genetic diversity decreased with the increase in distance from the center. Another region, in northern Hebei Province（115°–120°E×40°–42.5°N）, was observed having higher diversity than any surrounding regions, indicating that this is a sub-center of soybean diversity. Based on the presented results, the domestication and origin of soybean are also discussed.
基金supported by the Program of Accurate Identification and Display of Soybean Germplasm, China (NB08-2130315-(25-31)-06, NB07-2130315-(25-30)-06, NB06-070401-(22-27)-05), NB2010-2130315-25-05)the National Crop Germplasm Platform, Ministry of Agriculture and Ministry of Science and Technology, China (2012-004, 2014-004)
文摘The growth periods（GPs, from planting/emergence to reproductive stage 8（R8） of soybean cultivars vary in different ecological regions, especially in China with a very complex soybean cropping system. In this study, a 3-yr experimental study was undertaken in three geographical locations of China from 2008 to 2010, including the Northeast（40.66-45.85°N）, Huang-Huai（34.75-38.04°N） and southern（22.82-30.60°N） eco-regions with about 250 accessions in each region to clarify the classification of maturity group（MG） and identify the cultivars with stable GP to increase the knowledge about the GP distribution of soybean cultivars in China. GPs of soybean cultivars in different eco-regions were significant different with a gradual decrease from 115-125 d in the Northeast part to the 85-100 d in the southern part of China. The geographical location was the major factor for GP of cultivars from the Northeast, while the year of planting was the major factor affecting the stability of GPs in Huang-Huai summer and southern summer soybean. AMMI2（additive main effects and multiplicativeinteraction）-Biplot analysis showed that the GPs of soybean cultivars from the Northeast eco-region have a comparatively satisfactory environmental stability. Moreover, soybean cultivars with moderate GP/MG and stable environment adaptability in different eco-regions were identified based on the linear regression and AMMI analysis, which was important for the accurate classification of soybean MGs in future. Taken together, our results reflected the genetic diversity, geographical distribution and environmental stability of the Chinese soybean GP trait. Soybean cultivars with stable GP for various Chinese eco-regions would be beneficial for Chinese soybean genetic improvement, varietal introduction, exchange, and soybean breeding program for wide adaptability.
基金supported by the Platform of National Crop Germplasm Recourses of China and the Agricultural Science and Technology Innovation Program (ASTIP) of Chinese Academy of Agricultural Sciences
文摘The leaf is the main photosynthetic organ of plants,and it plays a significant role in the yield of crop species.Identifying the causal mutations and candidate genes that underlie leaf phenotypic variation is an important breeding target in soybean grain yield improvement.An ethyl methyl sulfonate(EMS)-induced soybean mutant DWARFCRINKLEDLEAF1(DCL1)with an aberrant crinkled leaf phenotype was identified in the background of the soybean cultivar Zhongpin 661(Zp661).We constructed an F2 segregating population from a cross between Zp661 and DCL1 in order to investigate the genomic locus associated with the crinkled leaf trait.Using bulk segregant analysis(BSA)combined with the whole-genome resequencing method,the Euclidean distance(ED)correlation algorithm detected 12 candidate genomic regions with a total length of 20.32 Mb that were linked to the target trait.Following a comparative analysis of the sequence data for the wild-type and mutant pools,only one single nucleotide mutation(C:G>T:A)located on the first exon of Glyma.19G207100 was found to be associated with the trait.Candidate gene validation based on a CAPS marker derived from the detected single-nucleotide polymorphism(SNP)indicated a nucleotide polymorphism between the two parents.Therefore,our findings reveal that Glyma.19G207100,which is renamed as GLYCINE MAX DWARF CRINKLED LEAF 1(GmDCL1),is a promising candidate gene involved in the morphogenesis of the crinkled leaf trait of the soybean mutant DCL1.This study provides a basis for the functional validation of this gene,with prospects for soybean breeding targeting grain yield enhancement.
基金supported by the National Basic Research Program of China (973 Program,2010CB125900 and 2004CB117203)the National Key Technologies R&D Program during the 11th Five-Year-Plan period (2006BAD13B05)+1 种基金the National Natural Science Foundation of China (30490251 and30471096)the Academy and Institute Foundation for Basic Scientific Research in Institute of Crop Sciences,Chinese Academy of Agricultural Sciences
文摘For clarifying the hierarchical patterns of population structure of soybean landraces in China, the seven clusters previously identified using Bayesian clustering of 1 504 soybean landraces based on SSR markers genotyping data were further analyzed. Using the largest value of AK, these landraces could be split into 20 sub-clusters, which was supported by highly significant pairwise Fst-values and generally in accordance with the geographic origin and sowing types. The autumn-sowing types ended up in one distinct sub-cluster from the otherwise summer-sowing type, where the autumn- sowing types are most likely derived from. The division into 20 sub-clusters explained 7.3% of the genetic variation, next to 9.7% present among the seven clusters, 81.1% residing among landraces within sub-clusters, and 1.9% within the landraces. The distribution pattern of genetic diversity among the sub-clusters of each cluster was uneven, with two HSuM sub-clusters （Central China） and some South China sub-clusters showing significantly higher level of genetic diversity.
