油料作物合辑Oil Crops

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Overexpression of Brassica napus cytosolic fructose-1,6-bisphosphatase and sedoheptulose-1,7-bisphosphatase genes significantly enhanced tobacco growth and biomass

LI Yan-yan, GUO Li-na, LIANG Cheng-zhen, MENG Zhi-gang, Syed Tahira, GUO San-dui, ZHANG Rui

2022, 21 (1): 49-59. DOI: 10.1016/S2095-3119(20)63438-4

Abstract （110） PDF in ScienceDirect

Elevated activities of cytosolic fructose-1,6-bisphosphatase (cyFBPase) and sedoheptulose-1,7-bisphosphatase (SBPase) are associated with higher yields in plants. In this study, the expression levels of the cyFBPase and SBPase genes were increased by overexpressing rape (Brassica napus) cDNA in tobacco (Nicotiana tabacum) plants. The transgenic plants co-expressing cyFBPase and SBPase (TpFS), or expressing single cyFBPase (TpF) or SBPase (TpS) had 1.77-, 1.55-, 1.23-fold cyFBPase and 1.45-, 1.12-, 1.36-fold SBPase activities as compared to the wild-type (WT), respectively. Photosynthesis rates of TpF, TpS and TpFS increased 4, 20 and 25% compared with WT plants. The SBPase and cyFBPase positively regulated each other and functioned synergistically in transgenic tobacco plants. In addition, the sucrose contents of the three transgenic plants were higher than that of WT plants. The starch accumulation of the TpFS and TpS plants was improved by 53 and 37%, but slightly decreased in TpF plants. Moreover, the transgenic tobacco plants harbouring SBPase and/or cyFBPase genes showed improvements in their growth, biomass, dry weight, plant height, stem diameter, leaf size, flower number, and pod weight. In conclusion, co-expression of SBPase and cyFBPase may pave a new way for improving crop yield in agricultural applications.

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Transcriptional profiling between yellow- and black-seeded Brassica napus reveals molecular modulations on flavonoid and fatty acid content

RONG Hao, YANG Wen-jing, XIE Tao, WANG Yue, WANG Xia-qin, JIANG Jin-jin, WANG You-ping

2022, 21 (8): 2211-2226. DOI: 10.1016/S2095-3119(21)63656-0

Abstract （157） PDF in ScienceDirect

Brassica napus is an important cash crop broadly grown for the vegetable and oil values. Yellow-seeded B. napus is preferred by breeders due to its improved oil and protein quality, less pigments and lignin compared with the black-seeded counterpart. This study compared the differences in flavonoid and fatty acid contents between yellow rapeseed from the progenies of B. napus–Sinapis alba somatic hybrids and the black-seeded counterpart using RNA-seq analysis. Through HPLC-PDA-ESI(−)/MS² analysis, it was found that phenylpropanoids and flavonoids (i.e., isorhamnetin, epicatechin, kaempferol, and other derivatives) in yellow seed were significantly lower than those in black seed. The fatty acid (FA) content in yellow rapeseed was higher than that in black rapeseed due to the variation of C16:0, C18:0, C18:1, C18:2, and C18:3 contents. RNA-seq analysis of seeds at four and five weeks after flowering (WAF) indicated that differentially expressed genes (DEGs) between black and yellow rapeseeds were enriched in flavonoid and FA biosynthesis, including BnTT3, BnTT4, BnTT18, and BnFAD2. Also, genes related to FA biosynthesis, desaturation and elongation (FAD3, LEC1, FUS3, and LPAT2) in yellow seed were up-regulated compared to those in black seed, while genes involved in beta-oxidation cycle (AIM1 and KAT2) of yellow seed were down-regulated compared to those in black seed. The DEGs related to the variation of flavonoids, phenylpropanoids, and FAs would help improve the knowledge of yellow seed character in B. napus and promote rapeseed improvement.

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Exogenous strigolactones promote lateral root growth by reducing the endogenous auxin level in rapeseed

MA Ni, WAN Lin, ZHAO Wei, LIU Hong-fang, LI Jun, ZHANG Chun-lei

2020, 19 (2): 465-482. DOI: 10.1016/S2095-3119(19)62810-8

Abstract （106） PDF in ScienceDirect

Strigolactones (SLs) are newly discovered plant hormones which regulate the normal development of different plant organs, especially root architecture. Lateral root formation of rapeseed seedlings before winter has great effects on the plant growth and seed yield. Here, we treated the seedlings of Zhongshuang 11 (ZS11), an elite conventional rapeseed cultivar, with different concentrations of GR24 (a synthetic analogue of strigolactones), and found that a low concentration (0.18 µmol L^–1) of GR24 could significantly increase the lateral root growth, shoot growth, and root/shoot ratio of seedlings. RNA-Seq analysis of lateral roots at 12 h, 1 d, 4 d, and 7 d after GR24 treatment showed that 2 301, 4 626, 1 595, and 783 genes were significantly differentially expressed, respectively. Function enrichment analysis revealed that the plant hormone transduction pathway, tryptophan metabolism, and the phenylpropanoid biosynthesis pathway were over-represented. Moreover, transcription factors, including AP2/ERF, AUX/IAA, NAC, MYB, and WRKY, were up-regulated at 1 d after GR24 treatment. Metabolomics profiling further demonstrated that the amounts of various metabolites, such as indole-3-acetic acid (IAA) and cis-zeatin were drastically altered. In particular, the concentrations of endogenous IAA significantly decreased by 52.4 and 75.8% at 12 h and 1 d after GR24 treatment, respectively. Our study indicated that low concentrations of exogenous SLs could promote the lateral root growth of rapeseed through interaction with other phytohormones, which provides useful clues for the effects of SLs on root architecture and crop productivity.

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Optimizing agronomic practices for closing rapeseed yield gaps under intensive cropping systems in China

ZHANG Zhi, CONG Ri-huan, REN Tao, LI Hui, ZHU Yun, LU Jian-wei

2020, 19 (5): 1241-1249. DOI: 10.1016/S2095-3119(19)62748-6

Abstract （84） PDF in ScienceDirect

A yield gap analysis for rapeseed (Brassica napus L.) is critical to meeting the oil demand by identifying yield potential and yield constraints. In this study, potential yield (Y_p), attainable yield (Y_att), and actual yield (Y_act) for winter rapeseed were determined in five different zones of China. A boundary line approach was adopted to calculate Yp, based on a large-scale field experimental database. A meta-analysis was conducted on the data obtained from 118 published studies to evaluate the effects of agronomic factors on rapeseed yield. The main results indicated that farmers only achieved 37–56% of the yield potential across the zones. The low altitude areas (L-URY) and lower reaches (LRY) of the Yangtze River Basin (YRB), China had high yield levels. The total yield gap was 1 893 kg ha^–1, due to the agronomic management factors, environmental factors, and socioeconomic factors. The meta-analysis showed that weed control and drainage were the best management practices to improve yields (45.6 and 35.3%, respectively), and other practices improved yields by 17.1–21.6%. Consequently, to narrow the yield gap over the short term, the study could focus on techniques that are easily implemented to farmers.

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Increased BnaMFT-transcript level is associated with secondary dormancy in oilseed rape (Brassica napus L.)

LIU Lei, FAN Wen-qi, LIU Fu-xia, YI Xin, TANG Tang, ZHOU Ying, TANG Zi-wei, CHEN Gui-min, ZHAO Xiang-xiang

2020, 19 (6): 1565-1576. DOI: 10.1016/S2095-3119(19)62684-5

Abstract （85） PDF in ScienceDirect

Brassica napus cultivars have little or no primary dormancy; however, they are prone to secondary dormancy induction. Secondary dormant seeds can produce volunteer plants, which can result in genetic contamination, reduced quality and biosafety issues. However, information regarding the molecular mechanism underlying secondary dormancy is limited. The MOTHER OF FT AND TFL1 (MFT) gene, which is evolutionarily conserved in the plant kingdom, acts in a complex gene network in the seed dormancy or germination processes. In this study, we identified four B. napus genes that share high homology with AtMFT, named as BnaMFT. Analyses of cis-acting elements showed that BnaMFT promoters contain multiple seed-specific regulatory elements, and various stress- and hormone-responsive elements. Further experiments validated that BnaMFTs were specifically expressed during seed maturation and in the dry seed, with peaks at 35–42 days after pollination. BnaMFTs were not sufficient for primary dormancy; however, they were significantly enhanced by secondary dormancy induction with PEG6000 treatment. Moreover, BnaMFT transcripts were elevated by treatment with abscisic acid (ABA), which is known to be accumulated during secondary dormancy. These results collectively suggest that increased BnaMFT transcription levels are associated with secondary dormancy induction in an ABA-dependent manner in B. napus.

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Differentially expressed miRNAs in anthers may contribute to the fertility of a novel Brassica napus genic male sterile line CN12A

Dong Yun, Wang Yi, Jin Feng-wei, Xing Li-juan, Fang Yan, Zhang Zheng-ying, ZOU Jun-jie, Wang Lei, Xu Miao-yun

2020, 19 (7): 1731-1742. DOI: 10.1016/S2095-3119(19)62780-2

Abstract （69） PDF in ScienceDirect

In Brassica napus L. (rapeseed), complete genic male sterility (GMS) plays an important role in the utilization of heterosis. Although microRNAs (miRNAs) play essential regulatory roles during bud development, knowledge of how GMS is regulated by miRNAs in rapeseed is rather limited. In this study, we obtained a novel recessive GMS system, CN12AB. The sterile line CN12A has defects in tapetal differentiation and degradation. Illumina sequencing was employed to examine the expression of miRNAs in the buds of CN12A and the fertile line CN12B. We identified 85 known miRNAs and 120 novel miRNAs that were expressed during rapeseed anther development. When comparing the expression levels of miRNAs between CN12A and CN12B, 19 and 18 known miRNAs were found to be differentially expressed in 0.5–1.0 mm buds and in 2.5–3.0 mm buds, respectively. Among these, the expression levels of 14 miRNAs were higher and the levels of 23 miRNAs were lower in CN12A compared with CN12B. The predicted target genes of these differentially expressed miRNAs encode protein kinases, F-box domain-containing proteins, MADS-box family proteins, SBP-box gene family members, HD-ZIP proteins, floral homeotic protein APETALA 2 (AP2), and nuclear factor Y, subunit A. These targets have previously been reported to be involved in pollen development and male sterility, suggesting that miRNAs might act as regulators of GMS in rapeseed anthers. Furthermore, RT-qPCR data suggest that one of the differentially expressed miRNAs, bna-miR159, plays a role in tapetal differentiation by regulating the expression of transcription factor BnMYB101 and participates in tapetal degradation and influences callose degradation by manipulating the expression of BnA6. These findings contribute to our understanding of the roles of miRNAs during anther development and the occurrence of GMS in rapeseed.

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Unraveling waterlogging tolerance-related traits with QTL analysis in reciprocal intervarietal introgression lines using genotyping by sequencing in rapeseed (Brassica napus L.)

DING Xiao-yu, XU Jin-song, HUANG He, QIAO Xing, SHEN Ming-zhen, CHENG Yong, ZHANG Xue-kun

2020, 19 (8): 1974-1983. DOI: 10.1016/S2095-3119(19)62783-8

Abstract （86） PDF in ScienceDirect

Soil waterlogging is a major environmental stress that suppresses the growth and productivity of rapeseed (Brassica napus L.). Natural genetic variations in waterlogging tolerance (WT) were observed but no QTL mapping has been done for WT related traits in rapeseed. In this study, QTL associated with three WT related traits including relative root length (RRL), relative hypocotyl length (RHL) and relative fresh weight (RFW) were dissected using a set of reciprocal introgression lines (ILs) derived from the cross GH01×ZS9, which showed significant difference in WT. Genotyping-by-sequencing (GBS) of the populations were performed, totally 1 468 and 1 450 binned SNPs were identified for GIL (GH01 as the recurrent parent) and ZIL (ZS9 as the recurrent parent) population, respectively. A total of 66 distinct QTLs for WT at the seedling establishment stage including 31 for RRL, 17 for RHL and 18 for RFW were detected. Among the 66 QTLs, 20 (29.4%) QTLs were detected in both genetic backgrounds and then they were integrated into six QTL clusters, which can be targeted in rapeseed breeding for improvement of WT through marker-assisted selection (MAS). Based on the physical positions of SNPs and the functional annotation of the Arabidopsis thaliana genome, 56 genes within the six QTL cluster regions were selected as preliminary candidate genes, then the resequencing and transcriptome information about parents were applied to narrow the extent of candidate genes. Twelve genes were determined as candidates for the six QTL clusters, some of them involved in RNA/protein degradation, most of them involved in oxidation-reduction process. These findings provided genetic resources, candidate genes to address the urgent demand of improving WT in rapeseed breeding.

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Crop photosynthetic response to light quality and light intensity

Iram SHAFIQ, Sajad HUSSAIN, Muhammad Ali RAZA, Nasir IQBAL, Muhammad Ahsan ASGHAR, Ali RAZA, FAN Yuan-fang, Maryam MUMTAZ, Muhammad SHOAIB, Muhammad ANSAR, Abdul MANAF, YANG Wen-yu, YANG Feng

2021, 20 (1): 4-23. DOI: 10.1016/S2095-3119(20)63227-0

Abstract （120） PDF in ScienceDirect

Under natural conditions, plants constantly encounter various biotic and abiotic factors, which can potentially restrict plant growth and development and even limit crop productivity. Among various abiotic factors affecting plant photosynthesis, light serves as an important factor that drives carbon metabolism in plants and supports life on earth. The two components of light (light quality and light intensity) greatly affect plant photosynthesis and other plant’s morphological, physiological and biochemical parameters. The response of plants to different spectral radiations and intensities differs in various species and also depends on growing conditions. To date, much research has been conducted regarding how different spectral radiations of varying intensity can affect plant growth and development. This review is an effort to briefly summarize the available information on the effects of light components on various plant parameters such as stem and leaf morphology and anatomy, stomatal development, photosynthetic apparatus, pigment composition, reactive oxygen species (ROS) production, antioxidants, and hormone production.

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Development and characterization of new allohexaploid resistant to web blotch in peanut

WANG Si-yu, LI Li-na, FU Liu-yang, LIU Hua, QIN Li, CUI Cai-hong, MIAO Li-juan, ZHANG Zhong-xin, GAO Wei, DONG Wen-zhao, HUANG Bing-yan, ZHENG Zheng, TANG Feng-shou, ZHANG Xin-you, DU Pei

2021, 20 (1): 55-64. DOI: 10.1016/S2095-3119(20)63228-2

Abstract （67） PDF in ScienceDirect

Peanut diseases seriously threaten peanut production, creating disease-resistant materials via interspecific hybridization is an effective way to deal with this problem. In this study, the embryo of an interspecific F₁ hybrid was obtained by crossing the Silihong (Slh) cultivar with Arachis duranensis (ZW55), a diploid wild species. Seedlings were generated by embryo rescue and tissue culture. A true interspecific hybrid was then confirmed by cytological methods and molecular markers. After treating seedlings with colchicine during in vitro multiplication, the established interspecific F₁ hybrid produced seeds which were named as Am1210. With oligonucleotide fluorescence in situ hybridization (Oligo FISH), molecular marker evaluations, morphological and web blotch resistance characterization, we found that: 1) Am1210 was an allohexaploid between Slh and ZW55; 2) the traits of spreading lateral branches, single-seeded or double-seeded pods and red seed coats were observed to be dominant compared to the erect type, multiple-seeded pods and brown seed coats; 3) the web blotch resistance of Am1210 was significantly improved than that of Slh, indicating the contribution of the web blotch resistance from the wild parent A. duranensis. In addition, 69 dominant and co-dominant molecular markers were developed which could be both used to verify the hybrid in this study and to identify translocation or introgression lines with A. duranensis chromosome fragments in future studies as well.

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Fine mapping and genetic analysis of resistance genes, Rsc18, against soybean mosaic virus

LIU Sang-lin, CHENG Yan-bo, MA Qi-bin, LI Mu JIANG Ze, XIA Qiu-ju, NIAN Hai

2022, 21 (3): 644-653. DOI: 10.1016/S2095-3119(20)63569-9

Abstract （150） PDF in ScienceDirect

Soybean mosaic virus (SMV) affects seed quality and production of soybean (Glycine max (L.) Merr.) worldwide. SC18 is one of the dominant SMV strains in South China, and accession Zhonghuang 24 displayed resistance to SC18. The F₁, F₂ and 168 F₁₁ recombinant inbred lines (RILs) population derived from a hybridization between Zhonghuang 24 (resistant, R) and Huaxia 3 (susceptible, S) were used in this study. According to the segregation ratios of the F₂ generation (3R:1S) and the recombinant inbred lines (RILs) population (1R:1S), one dominant locus may regulate the resistance to SC18 in Zhonghuang 24. By using composite interval mapping (CIM), Rsc18 was mapped to a 415.357-kb region on chromosome 13. Three candidate genes, including one NBS-LRR type gene and two serine/threonine protein type genes, were identified according to the genetic annotations, which may be related to the resistance to SC18. The qRT-PCR demonstrated that these genes were up-regulated in the R genotype compared to the control. In conclusion, the findings of this research enhanced the understanding about the R genes at the Rsc18 locus. Moreover, our results will provide insights for designing molecular markers to improve marker-assisted selection and developing new varieties with resistance to SC18.

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Image-based root phenotyping for field-grown crops: An example under maize/soybean intercropping

HUI Fang, XIE Zi-wen, LI Hai-gang, GUO Yan, LI Bao-guo, LIU Yun-ling, MA Yun-tao

2022, 21 (6): 1606-1619. DOI: 10.1016/S2095-3119(20)63571-7

Abstract （166） PDF in ScienceDirect

Root architecture, which determines the water and nutrient uptake ability of crops, is highly plastic in response to soil environmental changes and different cultivation patterns. Root phenotyping for field-grown crops, especially topological trait extraction, is rarely performed. In this study, an image-based semi-automatic root phenotyping method for field-grown crops was developed. The method consisted of image acquisition, image denoising and segmentation, trait extraction and data analysis. Five global traits and 40 local traits were extracted with this method. A good consistency in 1st-order lateral root branching was observed between the visually counted values and the values extracted using the developed method, with R²=0.97. Using the method, we found that the interspecific advantages for maize mainly occurred within 5 cm from the root base in the nodal roots of the 5th–7th nodes, and that the obvious inhibition of soybean was mostly reflected within 20 cm from the root base. Our study provides a novel approach with high-throughput and high-accuracy for field research on root morphology and branching features. It could be applied to the 3D reconstruction of field-grown root system architecture to improve the inputs to data-driven models (e.g., OpenSimRoot) that simulate root growth, solute transport and water uptake.

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Identification of candidate genes related to soluble sugar contents in soybean seeds using multiple genetic analyses

PAN Wen-jing, HAN Xue, HUANG Shi-yu, YU Jing-yao, ZHAO Ying, QU Ke-xin, ZHANG Ze-xin, YIN Zhen-gong, QI Hui-dong, YU Guo-long, ZHANG Yong, XIN Da-wei, ZHU Rong-sheng, LIU Chun-yan, WU Xiao-xia, JIANG Hong-wei, HU Zhen-bang, ZUO Yu-hu, CHEN Qing-shan, QI Zhao-ming

2022, 21 (7): 1886-1902. DOI: 10.1016/S2095-3119(21)63653-5

Abstract （195） PDF in ScienceDirect

Soluble sugar content in seeds is an important quality trait of soybean. In this study, 57 quantitative trait loci (QTLs) related to soluble sugar contents in soybean seeds were collected from databases and published papers. After meta-overview-collinearity integrated analysis to refine QTL intervals, eight consensus QTLs were identified. To further verify the consensus QTLs, a population of chromosome segment substitution lines (CSSLs) was analyzed. Two lines containing fragments covering the regions of consensus QTLs and the recurrent parent were selected: one line showed high soluble sugar contents associated with a consensus QTL fragment, and the other line showed low soluble sugar contents. Transcriptome sequencing was conducted for these two lines at the early, middle, and late stages of seed development, which identified 158, 109 and 329 differentially expressed genes, respectively. Based on the analyses of re-sequencing data of the CSSLs and the consensus QTL region, three candidate genes (Glyma.19G146800, Glyma.19G122500, and Glyma.19G128500) were identified in the genetic fragments introduced from wild soybean. Sequence comparisons between the two CSSL parents SN14 and ZYD00006 revealed a single nucleotide polymorphism (SNP) mutation in the coding sequence of Glyma.19G122500, causing a non-synonymous mutation in the amino acid sequence that affected the predicted protein structure. A Kompetitive allele-specific PCR (KASP) marker was developed based on this SNP and used to evaluate the CSSLs. These results lay the foundation for further research to identify genes related to soluble sugar contents in soybean seeds and for future soybean breeding.

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Identification and characterization of long-InDels through whole genome resequencing to facilitate fine-mapping of a QTL for plant height in soybean (Glycine max L. Merr.)

LIU Chen, TIAN Yu, LIU Zhang-xiong, GU Yong-zhe, ZHANG Bo, LI Ying-hui, NA Jie, QIU Li-juan

2022, 21 (7): 1903-1912. DOI: 10.1016/S2095-3119(21)63675-4

Abstract （176） PDF in ScienceDirect

With the development of sequencing technology, insertions-deletions (InDels) have been increasingly reported to be involved in the genetic deter mination of agronomical traits. However, most studies have focused on the identification and application of short-InDels (1–15 bp) for genetic analysis. The objective of this study was to deeply deploy long-InDels (>15 bp) for the genetic analysis of important agronomic traits in soybean. A total of 13 573 polymorphic long-InDels were identified between parents Zhongpin 03-5373 (ZP) and Zhonghuang 13 (ZH), which were unevenly distributed on 20 chromosomes of soybean, varying from 321 in Chr11 to 1 246 in Chr18. Consistent with the distribution pattern of annotated genes, the average density of long-InDels in arm regions was significantly higher than that in pericentromeric regions at the P=0.01 level. A total of 2 704 (19.9% of total) long-InDels were located in genic regions, including 319 large-effect long-InDels, which resulted in truncated or elongated protein sequences. A previously identified QTL (qPH16) underlying plant height was further analyzed, and it was found that 26 out of 35 (74.3%) long-InDel markers located in the qPH16 region showed clear polymorphisms between parents ZP and ZH. Seven markers, including three long-InDels and four previously reported SNP markers, were used to genotype 242 recombinant inbred lines derived from ZP×ZH. As a result, the qPH16 locus was narrowed from a 960-kb region to a 477.55-kb region, containing 65 annotated genes. Therefore, these long-InDels are a complementary genetic resource of SNPs and short-InDels for plant height and can facilitate genetic studies and molecular assisted selection breeding in soybean.

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Establishment and application of an accurate identification method for fragrant soybeans

ZHANG Yong-fang, ZHANG Chun-yan, ZHANG Bo, YIN Man, HONG Hui-long, YU Li-li, GAO Hua-wei, GU Yong-zhe, LIU Zhang-xiong, LI Fu-heng, QIU Li-juan

2021, 20 (5): 1193-1203. DOI: 10.1016/S2095-3119(20)63328-7

Abstract （98） PDF in ScienceDirect

In order to screen the aroma characteristics of soybean, a new method was established which can quickly quantify the content of 2-acetyl-1pyrroline (2-AP), an important compound related to soybean aroma, using gas chromatography-mass spectrometry (GC-MS). Based on peak profile, total peak area and retention time as test indexes, an accurate identification method for fragrant soybeans was established. The optimum parameters of the protocol consisted of column temperature 70°C, sample injector temperature 180°C, optimum extraction alcohol content 1 mL, NaCl content 0.1 g, ultrasonication time 10 min, and extraction time 1 h, which were established by using the orthogonal test of single factors and three factors with four levels (L₉(3)⁴). 2-AP content of leaves had significant correlations with seeds, which were easier to measure. The protocol was simple and easy to carry out, consumed only small amounts of reagents, and provided accurate and reliable results with good reproducibility. A total of 101 soybean genotypes from different geographical sources were analyzed using this protocol. The results showed that the average content of 2-AP was 0.29 mg L^–1, ranging from 0.094 to 1.816 mg L^–1, and the genetic diversity index was 0.54. Among all genotypes-tested, they were classified into three grades, including seven elite genotypes identified as “grade one fragrant soybeans”, which were Zhonglong 608, Heinong 88, Ha13-2958, Hongmiandou, Heinong 82, Huangmaodou, and Jiyu 21. These results provide both an identification technique and several elite aroma genotypes for gene discovery and good quality breeding in soybean.

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Ultrastructural studies of seed coat and cotyledon during rapeseed maturation

CAO Jian-bo, HE Li-min, Chinedu Charles NWAFOR, QIN Li-hong, ZHANG Chun-yu, SONG Yan-tun, HAO Rong

2021, 20 (5): 1239-1249. DOI: 10.1016/S2095-3119(20)63189-6

Abstract （78） PDF in ScienceDirect

Brassica napus L. (B. napus) is an important oil crop worldwide and it rapidly accumulates oil at late stage of seed maturation. However, little is known about the cellular mechanism of oil accumulation and seed color changes during the late stage of rapeseed development. Here, we analyzed the ultrastructure of seed coat, aleurone and cotyledon in embryos of B. napus from 25 to 70 days after flowering (DAF). The pigments, which were deposited on the cell wall of palisade cells in seed coat, determined dark black color of rapeseed. The chloroplasts degenerated into non-photosynthetic plastids which caused the green cotyledon to turn into yellow. The chloroplasts in aleurone and cotyledon cells respectively degenerated into remnants without inner and outer envelope membranes and ecoplasts with intact inner and outer envelope membranes. From 40 to 70 DAF, there were degraded chloroplasts without thylakoid, oil bodies contacting with plastids or protein bodies, big starch deposits of chloroplasts degrading into small particles then disappearing, and small endoplasmic reticulum (ER) in aleurone and cotyledon cells. Additionally, there were decreases of chlorophyll content and dramatic increases of oil content in rapeseed. These results suggested that the rapid oil accumulation was independent on the NADPH synthesized by photosynthesis of chloroplasts and probably utilized other sources of reductant, such as the oxidative pentose phosphate pathway during the late stage of rapeseed development. The triacylglycerol assembly presumably utilizes the enzymes in the plastid, cytosol or oil body of cotyledon and aleurone cells.

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Effects of shading stress during the reproductive stages on photosynthetic physiology and yield characteristics of peanut (Arachis hypogaea Linn.)

WANG Yi-bo, HUANG Rui-dong, ZHOU Yu-fei

2021, 20 (5): 1250-1265. DOI: 10.1016/S2095-3119(20)63442-6

Abstract （102） PDF in ScienceDirect

In intercropping systems, high-positioned crops often exert shading stress on low-positioned crops, thus affecting the agronomic characteristics of the low-positioned crops. This study determined the mechanisms of photosynthetic, physiological and yield variations among peanut cultivars under shading stress. Four peanut cultivars, S60, C4, P12, and YS151, were grown in the field and subjected to shading stress for 77 days during reproductive stages. S60 and P12 showed lower yield and reduced biomass accumulation than C4 and YS151 under shading stress. Shading stress induced higher antioxidant enzyme activities in S60 and P12, relative to C4 and YS151. Under shading stress, S60 and P12 showed a higher light-trapping capability than C4 and YS151, which was associated with changes in chlorophyll (Chl) a and b contents, and Chl a/b ratio. The net photosynthetic rate, stomatal conductance and transpiration rates of C4 and YS151 were decreased, but the intercellular CO₂ concentration increased under shading stress. The results demonstrated that non-stomatal limiting factors decreased the photosynthetic capacity of peanut under shading stress. The maximum photochemical efficiency of PSII (F_v/F_m) and non-photochemical quenching (NPQ) were higher in S60 and P12 than in C4 and YS151 under shading stress. These results suggest that S60 and P12 could absorb more light energy from weak light environments for photosynthesis than C4 and YS151 and dissipate the excess energy in the form of heat to improve their light protection ability. This study explains the inter-variety differences in shading stress tolerance in peanut and provides physiological parameters for guiding the selection of shade-tolerant cultivars.

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Characterization of the petiole length in soybean compact architecture mutant M657 and the breeding of new lines

GAO Hua-wei, SUN Ru-jian, YANG Meng-yuan, YAN Long, HU Xian-zhong, FU Guang-hui, HONG Hui-long, GUO Bing-fu, ZHANG Xiang, LIU Li-ke, ZHANG Shu-zhen, QIU Li-juan

2022, 21 (9): 2508-2520. DOI: 10.1016/j.jia.2022.07.004

Abstract （220） PDF in ScienceDirect

Phenotypic screening of soybean germplasm suitable for high planting density is currently the most viable strategy to increase yield. Previous studies have shown that soybean varieties with dwarf features and a short petiole often exhibit a compact plant architecture which could improve yield through increased planting density, although previously reported short petiole accessions were ultimately not usable for breeding in practice. Here, we established a method to assess petiole length and identified an elite mutant line, M657, that exhibits high photosynthetic efficiency. The agronomic traits of M657 were evaluated under field conditions, and appeared to be stable for short petiole across seven locations in northern, Huang–Huai, and southern China from 2017 to 2018. Compared with the Jihuang 13 wild type, the mutant M657 was shorter in both petiole length and plant height, exhibited lower total area of leaf, seed weight per plant and 100-seed weight, but had an increased number of effective branches and the growth period was prolonged by 2–7 days. Using M657 as a parental line for crosses with four other elite lines, we obtained four lines with desirable plant architecture and yield traits, thus demonstrating the feasibility of adopting M657 in breeding programs for soybean cultivars of high density and high yield.

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Genome-wide association and linkage mapping strategies reveal genetic loci and candidate genes of phosphorus utilization in soybean

ZHANG Hua, WU Hai-yan, TIAN Rui, KONG You-bin, CHU Jia-hao, XING Xin-zhu, DU Hui, JIN Yuan, LI Xi-huan, ZHANG Cai-ying

2022, 21 (9): 2521-2537. DOI: 10.1016/j.jia.2022.07.005

Abstract （203） PDF in ScienceDirect

Insufficient available phosphorus in soil has become an important limiting factor for the improvement of yield and quality in soybean. The mining of QTLs and candidate genes controlling soybean phosphorus utilization related traits is a necessary strategy to solve this problem. In this study, 11 phosphorus utilization related traits of a natural population of 281 typical soybean germplasms and a recombinant inbred line (RIL) population of 270 lines were evaluated under different phosphorus conditions at two critical stages: the four-leaf stage as the seedling critical stage was designated as the T₁ stage, and the six-leaf stage as the flowering critical stage was designated as the T₂ stage. In total, 200 single nucleotide polymorphism (SNP) loci associated with phosphorus utilization related traits were identified in the natural population, including 91 detected at the T₁ stage, and 109 detected at the T₂ stage. Among these SNP loci, one SNP cluster (s715611375, ss715611377, ss715611379 and ss715611380) on Gm12 was shown to be significantly associated with plant height under the low phosphorus condition at the T₁ stage, and the elite haplotype showed significantly greater plant height than the others. Meanwhile, one pleiotropic SNP cluster (ss715606501, ss715606506 and ss715606543) on Gm10 was found to be significantly associated with the ratio of root/shoot, root and total dry weights under the low phosphorus condition at the T₂ stage, and the elite haplotype also presented significantly higher values for related characteristics under the phosphorus starvation condition. Furthermore, four co-associated SNP loci (ss715597964, ss715607012, ss715622173 and ss715602331) were identified under the low phosphorus condition at both the T₁ and T₂ stages, and 12 QTLs were found to be consistent with these genetic loci in the RIL population. More importantly, 14 candidate genes, including MYB transcription factor, purple acid phosphatase, sugar transporter and HSP20-like chaperones superfamily genes, etc., showed differential expression levels after low phosphorus treatment, and three of them were further verified by qRT-PCR. Thus, these genetic loci and candidate genes could be applied in marker-assisted selection or map-based gene cloning for the genetic improvement of soybean phosphorus utilization.

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Linkage and association mapping of wild soybean (Glycine soja) seeds germinating under salt stress

SHI Mei-qi, LIAO Xi-liang, YE Qian, ZHANG Wei, LI Ya-kai, Javaid Akhter BHAT, KAN Gui-zhen, YU De-yue

2022, 21 (10): 2833-2847. DOI: 10.1016/j.jia.2022.07.031

Abstract （144） PDF in ScienceDirect

Salinity threatens soybean germination, growth and production. The germination stage is a key period in the life of soybean. Wild soybean contains many genes related to stress resistance that are valuable resources for the genetic improvement of soybean. To identify the genetic loci of wild soybean that are active during seed germination under salt stress, two populations, a soybean interspecific hybrid population comprising 142 lines and a natural population comprising 121 wild soybean accessions, were screened for three germination-related traits in this study. By using single-nucleotide polymorphism (SNP) markers with three salt tolerance indices, 25 quantitative trait loci (QTLs), 21 significant SNPs (–log₁₀(P)≥4.0) and 24 potential SNPs (3.5<–log₁₀(P)<4.0) were detected by linkage mapping and a genome-wide association study (GWAS) in two environments. The key genetic region was identified based on these SNPs and QTLs. According to the gene functional annotations of the W05 genome and salt-induced gene expression qRT-PCR analysis, GsAKR1 was selected as a candidate gene that responded to salt stress at the germination stage in the wild soybean. These results could contribute to determining the genetic networks of salt tolerance in wild soybean and will be helpful for molecular marker-assisted selection in the breeding of salt-tolerant soybean.

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Interactive effect of shade and PEG-induced osmotic stress on physiological responses of soybean seedlings

Muhammad Ahsan ASGHAR, JIANG Heng-ke, SHUI Zhao-wei, CAO Xi-yu, HUANG Xi-yu, Shakeel IMRAN, Bushra AHMAD, ZHANG Hao, YANG Yue-ning, SHANG Jing, YANG Hui, YU Liang, LIU Chun-yan, YANG Wen-yu, SUN Xin, DU Jun-bo

2021, 20 (9): 2382-2394. DOI: 10.1016/S2095-3119(20)63383-4

Abstract （94） PDF in ScienceDirect

Intensively farmed crops used to experience numerous environmental stresses. Among these, shade and drought significantly influence the morpho-physiological and biochemical attributes of plants. However, the interactive effect of shade and drought on the growth and development of soybean under dense cropping systems has not been reported yet. This study investigated the interactive effect of PEG-induced osmotic stress and shade on soybean seedlings. The soybean cultivar viz., C-103 was subjected to PEG-induced osmotic stress from polyethylene glycol 6000 (PEG-6000) under shading and non-shading conditions. PEG-induced osmotic stress significantly reduced the relative water contents, morphological parameters, carbohydrates and chlorophyll contents under both light environments. A significant increase was observed in osmoprotectants, reactive oxygen species and antioxidant enzymes in soybean seedlings. Henceforth, the findings revealed that, seedlings grown under non-shading conditions produced more malondialdehyde and hydrogen peroxide contents as compared to the shade-treated plants when subjected to PEG-induced osmotic stress. Likewise, the shaded plants accumulated more sugars and proline than non-shaded ones under drought stress. Moreover, it was found that non-shaded grown plants were more sensitive to PEG-induced osmotic stress than those exposed to shading conditions, which suggested that shade could boost the protective mechanisms against osmotic stress or at least would not exaggerate the adverse effects of PEG-induced osmotic stress in soybean seedlings.

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Identification and validation of stable and novel quantitative trait loci for pod shattering in soybean [Glycine max (L.) Merr.]

JIA Jia, WANG Huan, CAI Zhan-dong, WEI Ru-qian, HUANG Jing-hua, XIA Qiu-ju, XIAO Xiao-hui, MA Qi-bin, NIAN Hai, CHENG Yan-bo

2022, 21 (11): 3169-3184. DOI: 10.1016/j.jia.2022.08.082

Abstract （141） PDF in ScienceDirect

Pod shattering is an important domesticated trait which can cause great economic loss of crop yield in cultivated soybean. In this study, we utilized two recombinant inbred line populations (RILs, CY, Huachun 2×Wayao; GB, Guizao 1×B13) to identify quantitative trait loci (QTLs) associated with pod shattering in soybean across multiple environments. A total of 14 QTLs for pod shattering were identified in the two RIL populations, which had LOD scores ranging from 2.64 to 44.33 with phenotypic variance explanation (PVE) ranging from 1.33 to 50.85%. One QTL qPS16-1, located on chromosome 16, included a well-known functional gene Pod dehiscence 1 (Pdh1) that was reported previously. Ten new putative QTLs were validated in two RIL populations, and their LOD scores were between 2.55 and 4.24, explaining 1.33 to 2.60% of the phenotypic variation. Of which four novel QTLs (qPS01-1, qPS03-2, qPS05-1, and qPS07-1) could be detected in two environments where nine genes had specific changes in gene expression. Although the nine genes may have significant effects on pod shattering of soybean, their detailed functions still need to be further explored in the future. The results of this study will facilitate a better understanding of the genetic basis of the pod shattering-resistant trait and benefit soybean molecular breeding for improving pod shattering resistance

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Effects of drought stress on root morphology and spatial distribution of soybean and adzuki bean

Hyen Chung CHUN, Sanghun LEE, Young Dae CHOI, Dong Hyeok GONG, Ki Youl JUNG

2021, 20 (10): 2639-2651. DOI: 10.1016/S2095-3119(20)63560-2

Abstract （111） PDF in ScienceDirect

Due to global climate change, Korea is facing severe droughts that affect the planting and early vegetative periods of upland crops. Soybean and adzuki bean are important legume crops in Korea, so it is critical to understand their adaptations to water stress. This study investigated the changes in root morphological properties in soybean and adzuki bean and quantified the findings using fractal analysis. The experiment was performed at the National Institute of Crop Science in Miryang, Korea. Soybeans and adzuki beans were planted in test boxes and grown for 30 days. The boxes were filled with bed soil with various soil moisture treatments. Root images were obtained and scanned every two days, and the root properties were characterized by root length, depth and surface area, number of roots, and fractal parameters (fractal dimension and lacunarity). Root depth, length and surface area and the number of roots increased in both crops as the soil moisture content increased. The fractal dimension and lacunarity values increased as the soil moisture content increased. These results indicated that the greater the soil moisture, the more heterogeneous the root structure. Correlation analysis of the morphological properties and fractal parameters indicated that soybean and adzuki bean had different root structure developments. Both soybean and adzuki bean were sensitive to the amount of soil moisture in the early vegetative stage. Soybean required a soil moisture content greater than 70% of the field capacity to develop a full root structure, while adzuki bean required 100% of the field capacity. These results would be useful in understanding the responses of soybean and adzuki bean to water stress and managing irrigation during cultivation.

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Identification of quantitative trait loci and candidate genes controlling seed pigments of rapeseed

ZHU Mei-chen, HU Ran, ZHAO Hui-yan, TANG Yun-shan, SHI Xiang-tian, JIANG Hai-yan, ZHANG Zhi-yuan, FU Fu-you, XU Xin-fu, TANG Zhang-lin, LIU Lie-zhao, LU Kun, LI Jia-na, QU Cun-min

2021, 20 (11): 2862-2879. DOI: 10.1016/S2095-3119(20)63377-9

Abstract （112） PDF in ScienceDirect

Rapeseed (Brassica napus L.) is an important source of edible vegetable oil and feed protein; however, seed pigments affect the quality of rapeseed oil and the feed value of the residue from oil pressing. Here, we used a population of rapeseed recombinant inbred lines (RILs) derived from the black-seeded male parent cultivar Zhongyou 821 and the yellow-seeded female parent line GH06 to map candidate genes controlling seed pigments in embryos and the seed coat. We detected 94 quantitative trait loci (QTLs) for seed pigments (44 for embryos and 50 for seed coat), distributed over 15 of the 19 rapeseed chromosomes. These included 28 QTLs for anthocyanidin content, explaining 2.41–44.66% of phenotypic variation; 24 QTLs for flavonoid content, explaining 2.41–20.26% of phenotypic variation; 16 QTLs for total phenol content, accounting for 2.74–23.68% of phenotypic variation; and 26 QTLs for melanin content, accounting for 2.37–24.82% of phenotypic variation, indicating that these traits are under multigenic control. Consensus regions on chromosomes A06, A09 and C08 were associated with multiple seed pigment traits, including 15, 19 and 10 QTLs, respectively, most of which were major QTLs explaining >10% of the phenotypic variation. Based on the annotation of the B. napus “Darmor-bzh” reference genome, 67 candidate genes were predicted from these consensus QTLs regions, and 12 candidate genes were identified as potentially involved in pigment accumulation by RNA-seq and qRT-PCR analysis. These preliminary results provide insight into the genetic architecture of pigment biosynthesis and lay a foundation for exploring the molecular mechanisms underlying seed coat color in B. napus.

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Identification of the genetic locus associated with the crinkled leaf phenotype in a soybean (Glycine max L.) mutant by BSA-Seq technology

OCHAR Kingsley, SU Bo-hong, ZHOU Ming-ming, LIU Zhang-xiong, GAO Hua-wei, SOBHI F. Lamlom, QIU Li-juan

2022, 21 (12): 3524-3539. DOI: 10.1016/j.jia.2022.08.095

Abstract （214） PDF in ScienceDirect

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.

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Sexual compatibility of transgenic soybean and different wild soybean populations

HU Yu-qi, SHENG Ze-wen, LIU Jin-yue, LIU Qi, QIANG Shen, SONG Xiao-ling, LIU Biao

2022, 21 (1): 36-48. DOI: 10.1016/S2095-3119(20)63385-8

Abstract （177） PDF in ScienceDirect

The introduction of genetically modified (GM) soybean into farming systems raises great concern that transgenes from GM soybean may flow to endemic wild soybean via pollen. This may increase the weediness of transgenic soybean by increasing the fitness of hybrids under certain conditions and threaten the genetic diversity of wild soybean populations. Although pollen-mediated gene flow between GM crops and wild relatives is dependent on many factors, the sexual compatibility (SC) determined by their genetic backgrounds is the conclusive factor. The considerable genetic variation among wild soybean populations may cause compatibility differences between different wild and cultivated soybeans. Thus, an evaluation of the SC between transgenic soybean and different wild soybeans is essential for assessing the environmental consequences of cultivated soybean–wild soybean transgene flow. The podding and seed sets were assessed after artificial hybridization using transgenic glyphosate-resistant soybean as the paternal parent and 18 wild soybean populations as the maternal parents. Then, the average number of filled seeds produced in 200 flowers (AFS) was calculated for each wild soybean under natural self-pollination as well as under artificial crossing with transgenic soybean. Finally, the index of cross-SC was calculated (ICSC) as the ratio of the AFS of wild soybean artificially crossed with transgenic soybean and the AFS of naturally self-pollinated wild soybean. The results demonstrated that after self-pollination and crossing with transgenic soybean, the average podding rates of 18 wild soybean populations ranged within 96.50–99.50% and 4.92–18.03%, and the average filled seed numbers per pod varied from 1.70 to 2.69 and 0.20 to 0.48, respectively. The results showed that approximately 89% of wild soybeans displayed either medium or higher than medium SC with transgenic soybean (ICSC>1.0%). This implied the high possibility of gene flow via pollen from transgenic soybean to wild soybean.

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Co-silencing E1 and its homologs in an extremely late-maturing soybean cultivar confers super-early maturity and adaptation to high-latitude short-season regions

LIU Li-feng, GAO Le, ZHANG Li-xin, CAI Yu-peng, SONG Wen-wen, CHEN Li, YUAN Shan, WU Ting-ting, JIANG Bing-jun, SUN Shi, WU Cun-xiang, HOU Wen-sheng, HAN Tian-fu

2022, 21 (2): 326-335. DOI: 10.1016/S2095-3119(20)63391-3

Abstract （140） PDF in ScienceDirect

Soybean (Glycine max (L.) Merr.), a typical short-day plant, is sensitive to photoperiod, which limits the geographical range for its cultivation. In the flowering pathway regulated by photoperiod, E1, a flowering inhibitor in soybean, plays the dominant role in flowering time regulation. Two E1 homologs, E1-like-a (E1La) and E1-like-b (E1Lb), play overlapping or redundant roles in conjunction with E1. In the present study, E1 and E1La/b were simultaneously silenced via RNA interference (RNAi) in Zigongdongdou (ZGDD), an extremely late-flowering soybean landrace from southern China. As a result, RNAi lines showed a much earlier-flowering phenotype and obvious photoperiod insensitivity compared with wild-type (WT) plants. In RNAi transgenic plants, the expression levels of flowering inhibitor GmFT4 and flowering promoters GmFT2a/GmFT5a were significantly down- and up-regulated, respectively. Further, the maturity group (MG) of the RNAi lines was reduced from WT ZGDD’s MG VIII (extremely late-maturity) to MG 000 (super-early maturity), which can even grow in the northernmost village of China located at a latitude of 53.5°N. Our study confirms that E1 and E1La/b can negatively regulate flowering time in soybean. The RNAi lines generated in this study, with early flowering and maturity traits, can serve as valuable materials and a technical foundation for breeding soybeans that are adapted to high-latitude short-season regions.

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Field mold stress induced catabolism of storage reserves in soybean seed and the resulting deterioration of seed quality in the field

DENG Jun-cai, LI Xiao-man, XIAO Xin-li, WU Hai-jun, YANG Cai-qiong, LONG Xi-yang, ZHANG Qi-hui, Nasir Iqbal, WANG Xiao-chun, YONG Tai-wen, DU Jun-bo, YANG Feng, LIU Wei-guo, ZHANG Jing, WU Xiao-ling, WU Yu-shan, YANG Wen-yu, LIU Jiang

2022, 21 (2): 336-350. DOI: 10.1016/S2095-3119(20)63594-8

Abstract （148） PDF in ScienceDirect

Excessive rainfall provides a favorable condition for field mold infection of plants, which triggers field mold (FM) stress. If FM stress occurs during the late maturation stage of soybean seed, it negatively affects seed yield and quality. To investigate the responses of soybean seed against FM stress and identify the underlying biochemical pathways involved, a greenhouse was equipped with an artificial rain producing system to allow the induction of mold growth on soybean seed. The induced quality changes and stress responses were revealed on the levels of both transcriptome and metabolome. The results showed that soybean seeds produced under FM stress conditions had an abnormal and inferior appearance, and also contained less storage reserves, such as protein and polysaccharide. Transcriptional analysis demonstrated that genes involved in amino acid metabolism, glycolysis, tricarboxylic acid, β-oxidation of fatty acids, and isoflavone biosynthesis were induced by FM stress. These results were supported by a multiple metabolic analysis which exhibited increases in the concentrations of a variety of amino acids, sugars, organic acids, and isoflavones, as well as reductions of several fatty acids. Reprogramming of these metabolic pathways mobilized and consumed stored protein, sugar and fatty acid reserves in the soybean seed in order to meet the energy and substrate demand on the defense system, but led to deterioration of seed quality. In general, FM stress induced catabolism of storage reserves and diminished the quality of soybean seed in the field. This study provides a more profound insight into seed deterioration caused by FM stress.

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Identification of tolerance to high density and lodging in short petiolate germplasm M657 and the effect of density on yield-related phenotypes of soybean

GAO Hua-wei, YANG Meng-yuan, YAN Long, HU Xian-zhong, HONG Hui-long, ZHANG Xiang, SUN Ru-jian, WANG Hao-rang, WANG Xiao-bo, LIU Li-ke, ZHANG Shu-zhen, QIU Li-juan

2023, 22 (2): 434-446. DOI: 10.1016/j.jia.2022.08.047

Abstract （167） PDF in ScienceDirect

Soybean yield has 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 avenue to improve yield. We compared the relationship between yield-related traits, lodging resistance, and petiole-associated phenotypes in the short petiole germplasm M657 with three control accessions over 2017-2018 in four locations of the Huang-Huai region. The results showed M657 exhibited stable and high tolerance to high planting density and resistance to lodging, especially at the highest density (8×10⁵ plants ha^-1). Regression analysis showed that shorter petiole length was significantly associated with increased lodging resistance. Yield analysis showed that M657 achieved higher yields under higher densities, especially in the north Huang-Huai region. There are markedly different responses to intra- and inter-row spacing designs among varieties in both lodging and yield related to location and density. Lodging was positively correlated with planting density, plant height, petiole length, and number of effective branches, and negatively correlated with stem diameter, seed number per plant, and seed weight per plant. The yield of soybean was increased by appropriately increasing planting density on the basis of current soybean varieties in the Huang-Huai region. This study provides a valuable new germplasm resource for introgression of compact architecture traits amenable to high yield in high density planting systems and establishes a high-yield model of soybean in the Huang-Huai region.

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Millet/peanut intercropping at a moderate N rate increases crop productivity and N use efficiency, as well as economic benefits, under rain-fed conditions

LIU Zhu, NAN Zhen-wu, LIN Song-ming, YU Hai-qiu, XIE Li-yong, MENG Wei-wei, ZHANG Zheng, WAN Shu-bo

2023, 22 (3): 738-751. DOI: 10.1016/j.jia.2022.08.078

Abstract （198） PDF in ScienceDirect

Cereal and legume intercropping has been widely adopted to increase crop productivity in sustainable farming systems worldwide. Among different intercropping combinations, millet and peanut intercropping can be adapted to most water-limited areas. However, there are few studies on the differences in yield characteristics and nitrogen use efficiency between millet/peanut intercropping and monocultures under different nitrogen (N) application rates. The objective of this study was to determine the yield advantages and economic benefits, as well as the appropriate N application rate, of millet/peanut intercropping. A two-year field experiment was conducted with three cropping patterns (monoculture millet, monoculture peanut and millet/peanut intercropping) and four N rates (0, 75, 150 and 225 kg ha⁻¹). The results showed that the land equivalent ratio (LER) and net effect (NE) of the intercropping system reached their highest levels at the N input of 150 kg ha⁻¹ in 2018 and 2019 (1.04 for LER, 0.347 Mg ha⁻¹ for NE, averaged across two years). Millet was the dominant crop in the intercropping system (aggressivity of millet and peanut (Amp)>0, competitive ratio of millet and peanut (CRmp)>1), and millet yields achieved their highest values at N inputs of 225 kg ha⁻¹ for monoculture and 150 kg ha⁻¹ for intercropping. NUE reached its highest levels with N inputs of 150 kg ha⁻¹ for all planting patterns over the two years. Intercropping combined with an N input of 150 kg ha⁻¹ achieved the highest net income of 2 791 USD ha⁻¹, with a benefit-cost ratio of 1.56, averaged over the two years. From the perspective of economics and agricultural sustainable development, millet/peanut intercropping at 150 kg N ha⁻¹ seems to be a promising alternative to millet or peanut monoculture.

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QTL analysis for plant height and fine mapping of two environmentally stable QTLs with major effects in soybean

TIAN Yu, YANG Lei, LU Hong-feng, ZHANG Bo, LI Yan-fei, LIU Chen, GE Tian-li, LIU Yu-lin, HAN Jia-nan, LI Ying-hui, QIU Li-juan

2022, 21 (4): 933-946. DOI: 10.1016/S2095-3119(21)63693-6

Abstract （177） PDF in ScienceDirect

Plant height is an important agronomic trait, which is governed by multiple genes with major or minor effects. Of numerous QTLs for plant height reported in soybean, most are in large genomic regions, which results in a still unknown molecular mechanism for plant height. Increasing the density of molecular markers in genetic maps will significantly improve the efficiency and accuracy of QTL mapping. This study constructed a high-density genetic map using 4 011 recombination bin markers developed from whole genome re-sequencing of 241 recombinant inbred lines (RILs) and their bi-parents, Zhonghuang 13 (ZH) and Zhongpin 03-5373 (ZP). The total genetic distance of this bin map was 3 139.15 cM, with an average interval of 0.78 cM between adjacent bin markers. Comparative genomic analysis indicated that this genetic map showed a high collinearity with the soybean reference genome. Based on this bin map, nine QTLs for plant height were detected across six environments, including three novel loci (qPH-b_11, qPH-b_17 and qPH-b_18). Of them, two environmentally stable QTLs qPH-b_13 and qPH-b_19-1 played a major role in plant height, which explained 10.56–32.7% of the phenotypic variance. They were fine-mapped to 440.12 and 237.06 kb region, covering 54 and 28 annotated genes, respectively. Via the function of homologous genes in Arabidopsis and expression analysis, two genes of them were preferentially predicted as candidate genes for further study.

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Characterization of chromosome segment substitution lines reveals candidate genes associated with the nodule number in soybean

ZOU Jia-nan, ZHANG Zhan-guo, KANG Qing-lin, YU Si-yang, WANG Jie-qi, CHEN Lin, LIU Yan-ru, MA Chao, ZHU Rong-sheng, ZHU Yong-xu, DONG Xiao-hui, JIANG Hong-wei, WU Xiao-xia, WANG Nan-nan, HU Zhen-bang, QI Zhao-ming, LIU Chun-yan, CHEN Qing-shan, XIN Da-wei, WANG Jin-hui

2022, 21 (8): 2197-2210. DOI: 10.1016/S2095-3119(21)63658-4

Abstract （127） PDF in ScienceDirect

Soybean is one of the most important food crops worldwide. Like other legumes, soybean can form symbiotic relationships with Rhizobium species. Nitrogen fixation of soybean via its symbiosis with Rhizobium is pivotal for sustainable agriculture. Type III effectors (T3Es) are essential regulators of the establishment of the symbiosis, and nodule number is a feature of nitrogen-affected nodulation. However, genes encoding T3Es at quantitative trait loci (QTLs) related to nodulation have rarely been identified. Chromosome segment substitution lines (CSSLs) have a common genetic background but only a few loci with heterogeneous genetic information; thus, they are suitable materials for identifying candidate genes at a target locus. In this study, a CSSL population was used to identify the QTLs related to nodule number in soybean. Single nucleotide polymorphism (SNP) markers and candidate genes within the QTLs interval were detected, and it was determined which genes showed differential expression between isolines. Four candidate genes (GmCDPK28, GmNAC1, GmbHLH, and GmERF5) linked to the SNPs were identified as being related to nodule traits and pivotal processes and pathways involved in symbiosis establishment. A candidate gene (GmERF5) encoding a transcription factor that may interact directly with the T3E NopAA was identified. The confirmed CSSLs with important segments and candidate genes identified in this study are valuable resources for further studies on the genetic network and T3Es involved in the signaling pathway that is essential for symbiosis establishment.

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