Phenoxyacetic acid enhances nodulation symbiosis during the rapid growth stage of soybean.

Root exudates are known signaling agents that influence legume root nodulation, but the molecular mechanisms for nonflavonoid molecules remain largely unexplored. The number of soybean root nodules during the initial growth phase shows substantial discrepancies at distinct developmental junctures. Using a combination of metabolomics analyses on root exudates and nodulation experiments, we identify a pivotal role for certain root exudates during the rapid growth phase in promoting nodulation. Phenoxyacetic acid (POA) was found to activate the expression of GmGA2ox10 and thereby facilitate rhizobial infection and the formation of infection threads. Furthermore, POA exerts regulatory control on the miR172c-NNC1 module to foster nodule primordia development and consequently increase nodule numbers. These findings collectively highlight the important role of POA in enhancing nodulation during the accelerated growth phase of soybeans.

Inferring Physical Properties of Symmetric States from the Fewest Copies.

Learning physical properties of high-dimensional states is crucial for developing quantum technologies but usually consumes an exceedingly large number of samples which are difficult to afford in practice. In this Letter, we use the methodology of quantum metrology to tackle this difficulty, proposing a strategy built upon entangled measurements for dramatically reducing sample complexity. The strategy, whose characteristic feature is symmetrization of observables, is powered by the exploration of symmetric structures of states which are ubiquitous in physics. It is provably optimal under some natural assumption, efficiently implementable in a variety of contexts, and capable of being incorporated into existing methods as a basic building block. We apply the strategy to different scenarios motivated by experiments, demonstrating exponential reductions in sample complexity.

Peptide REF1 is a local wound signal promoting plant regeneration.

Plants frequently encounter wounding and have evolved an extraordinary regenerative capacity to heal the wounds. However, the wound signal that triggers regenerative responses has not been identified. Here, through characterization of a tomato mutant defective in both wound-induced defense and regeneration, we demonstrate that in tomato, a plant elicitor peptide (Pep), REGENERATION FACTOR1 (REF1), acts as a systemin-independent local wound signal that primarily regulates local defense responses and regenerative responses in response to wounding. We further identified PEPR1/2 ORTHOLOG RECEPTOR-LIKE KINASE1 (PORK1) as the receptor perceiving REF1 signal for plant regeneration. REF1-PORK1-mediated signaling promotes regeneration via activating WOUND-INDUCED DEDIFFERENTIATION 1 (WIND1), a master regulator of wound-induced cellular reprogramming in plants. Thus, REF1-PORK1 signaling represents a conserved phytocytokine pathway to initiate, amplify, and stabilize a signaling cascade that orchestrates wound-triggered organ regeneration. Application of REF1 provides a simple method to boost the regeneration and transformation efficiency of recalcitrant crops.

A novel miR160a-GmARF16-GmMYC2 module determines soybean salt tolerance and adaptation.

Salt stress is a major challenge that has a negative impact on soybean growth and productivity. Therefore, it is important to understand the regulatory mechanism of salt response to ensure soybean yield under such conditions. In this study, we identified and characterized a miR160a-GmARF16-GmMYC2 module and its regulation during the salt-stress response in soybean. miR160a promotes salt tolerance by cleaving GmARF16 transcripts, members of the Auxin Response Factor (ARF) family, which negatively regulates salt tolerance. In turn, GmARF16 activates GmMYC2, encoding a bHLH transcription factor that reduces salinity tolerance by down-regulating proline biosynthesis. Genomic analysis among wild and cultivated soybean accessions identified four distinct GmARF16 haplotypes. Among them, the GmARF16H3 haplotype is preferentially enriched in localities with relatively saline soils, suggesting GmARF16H3 was artificially selected to improve salt tolerance. Our findings therefore provide insights into the molecular mechanisms underlying salt response in soybean and provide valuable genetic targets for the molecular breeding of salt tolerance.

Identification and mapping of a recessive allele, dt3, specifying semideterminate stem growth habit in soybean.

KEY MESSAGE: A locus, dt3, modulating semideterminancy in soybean, was discovered by a combination of genome-wide association studies and linkage mapping with multiple distinct biparental populations. Stem growth habit is a key architectural trait in many plants that contributes to plant productivity and environmental adaptation. In soybean, stem growth habit is classified as indeterminate, semideterminate, or determinate, of which semideterminacy is often considered as a counterpart of the "Green Revolution" trait in cereals that significantly increased grain yields. It has been demonstrated that semideterminacy in soybean is modulated by epistatic interaction between two loci, Dt1 on chromosome 19 and Dt2 on chromosome 18, with the latter as a negative regulator of the former. Here, we report the discovery of a third locus, Dt3, modulating soybean stem growth habit, which was delineated to a ~ 196-kb region on chromosome 10 by a combination of allelic and haplotypic analysis of the Dt1 and Dt2 loci in the USDA soybean Germplasm Collection, genome-wide association studies with three subsets of the collection, and linkage mapping with four biparental populations derived from crosses between one of two elite indeterminate cultivars and each of four semideterminate varieties possessing neither Dt2 nor dt1. These four semideterminate varieties are recessive mutants (i.e., dt3/dt3) in the Dt1/Dt1;dt2/dt2 background. As the semideterminacy modulated by the Dt2 allele has unfavorable pleotropic effects such as sensitivity to drought stress, dt3 may be an ideal alternative for use to develop semideterminate cultivars that are more resilient to such an environmental stress. This study enhances our understanding of the genetic factors underlying semideterminacy and enables more accurate marker-assisted selection for stem growth habit in soybean breeding.

Non-Hermitian Floquet Topological Matter-A Review.

The past few years have witnessed a surge of interest in non-Hermitian Floquet topological matter due to its exotic properties resulting from the interplay between driving fields and non-Hermiticity. The present review sums up our studies on non-Hermitian Floquet topological matter in one and two spatial dimensions. We first give a bird's-eye view of the literature for clarifying the physical significance of non-Hermitian Floquet systems. We then introduce, in a pedagogical manner, a number of useful tools tailored for the study of non-Hermitian Floquet systems and their topological properties. With the aid of these tools, we present typical examples of non-Hermitian Floquet topological insulators, superconductors, and quasicrystals, with a focus on their topological invariants, bulk-edge correspondences, non-Hermitian skin effects, dynamical properties, and localization transitions. We conclude this review by summarizing our main findings and presenting our vision of future directions.

The conserved evolution of plant H+-ATPase family and the involvement of soybean H+-ATPases in sodium bicarbonate stress responses.

Plant plasma membrane (PM) H+-ATPases are essential pumps involved in multiple physiological processes. They play a significant role in regulating pH homeostasis and membrane potential by generating the electrochemical gradient of the proton across the plasma membrane. However, information on soybean PM H+-ATPase is still limited. In this study, we conducted the evolutionary analysis of PM H+-ATPases in land plants and investigated the subfamily classification and whole genome duplication of PM H+-ATPases in angiosperms. We further characterized the extremely high conservation of the soybean PM H+-ATPase family in terms of gene structure, domain architecture, and protein sequence identity. Using the yeast system, we confirmed the highly conserved biochemical characteristics (14-3-3 binding affinity and pump activity) of soybean PM H+-ATPases and their conserved function in enhancing tolerance to high pH and NaHCO3 stresses. Meanwhile, our results also revealed their divergence in the transcriptional expression in different tissues and under sodium bicarbonate stress. Finally, the function of soybean PM H+-ATPases in conferring sodium bicarbonate tolerance was validated using transgenic Arabidopsis. Together, these results conclude that the soybean PM H+-ATPase is evolutionarily conserved and positively regulates the response to sodium bicarbonate stress.

Effect of Heat Treatment on the Digestive Characteristics of Different Soybean Oil Body Emulsions.

Soybean oil body (SOB) emulsions were prepared using OBs extracted at pH 11.0 and pH 7.0. The pH 11.0-SOB comprised oleosins, whereas pH 7.0-SOB comprised extrinsic proteins and oleosins. All SOB emulsions were heated at 60-100 °C for 15 min. Heating may lead to the release of extrinsic proteins from the surface of pH 7.0-SOB due to heat-induced denaturation. The total proportion of α-helix and ß-sheets gradually decreased from 77 (unheated) to 36.2% (100 °C). During stomach digestion, the extrinsic protein hydrolysis of heated pH 7.0-SOB emulsions was fast between 60 and 80 °C, and it then slowed between 90 and 100 °C; heating inhibited the oleosin hydrolysis of pH 7.0- and 11.0-SOBs. Heat treatment promoted aggregation and coalescence, and it resulted in increased particle sizes for all emulsions. Larger aggregates were found in heated pH 7.0-SOB emulsions, and larger oil droplets were found in heated pH 11.0-SOB emulsions. After intestinal digestion, the droplets of all SOB emulsions gradually dispersed, and particle sizes decreased. Different heating temperatures had lesser effects on particle sizes and microstructures. Lipolysis was affected by the extraction pH and heating. For pH 11.0-SOB emulsions, the FFA release tendency was greatly affected by the heating temperature, and heating to 80 °C resulted in the highest FFA release (74%). However, all pH 7.0-SOB emulsions had similar total FFA releases. In addition, the droplet charges of heated pH 7.0-SOB emulsions were lower than those of unheated pH 7.0-SOB emulsions in both the intestine and stomach phases; however, the charge changes in different pH 11.0-SOB emulsions showed the opposite tendency. This study will offer guidance regarding the application of SOB emulsions in food.

A retrotransposon insertion in the Mao1 promoter results in erect pubescence and higher yield in soybean.

Adaptive changes in crops contribute to the diversity of agronomic traits, which directly or indirectly affect yield. The change of pubescence form from appressed to erect is a notable feature during soybean domestication. However, the biological significance and regulatory mechanism underlying this transformation remain largely unknown. Here, we identified a major-effect locus, PUBESCENCE FORM 1 (PF1), the upstream region of Mao1, that regulates pubescence form in soybean. The insertion of a Ty3/Gypsy retrotransposon in PF1 can recruit the transcription factor GAGA-binding protein to a GA-rich region, which up-regulates Mao1 expression, underpinning soybean pubescence evolution. Interestingly, the proportion of improved cultivars with erect pubescence increases gradually with increasing latitude, and erect-pubescence cultivars have a higher yield possibly through a higher photosynthetic rate and photosynthetic stability. These findings open an avenue for molecular breeding through either natural introgression or genome editing toward yield improvement and productivity.

Cold-induced calreticulin OsCRT3 conformational changes promote OsCIPK7 binding and temperature sensing in rice.

Unusually low temperatures caused by global climate change adversely affect rice production. Sensing cold to trigger signal network is a key base for improvement of chilling tolerance trait.  Here, we report that Oryza sativa Calreticulin 3 (OsCRT3) localized at the endoplasmic reticulum (ER) exhibits conformational changes under cold stress, thereby enhancing its interaction with CBL-interacting protein kinase 7 (OsCIPK7) to sense cold. Phenotypic analyses of OsCRT3 knock-out mutants and transgenic overexpression lines demonstrate that OsCRT3 is a positive regulator in chilling tolerance. OsCRT3 localizes at the ER and mediates increases in cytosolic calcium levels under cold stress. Notably, cold stress triggers secondary structural changes of OsCRT3 and enhances its binding affinity with OsCIPK7, which finally boosts its kinase activity. Moreover, Calcineurin B-like protein 7 (OsCBL7) and OsCBL8 interact with OsCIPK7 specifically on the plasma membrane. Taken together, our results thus identify a cold-sensing mechanism that simultaneously conveys cold-induced protein conformational change, enhances kinase activity, and Ca2+ signal generation to facilitate chilling tolerance in rice.

Digestive characteristics of oil body extracted from soybean aqueous extract at different pHs.

Soybean oil bodies (SOBs), a natural source of pre-emulsified oil, have important potential applications in food industry. In this study, SOBs were extracted from raw soybean milk at pH 5.0, 7.0, 8.0, 9.0 and 11.0. The pH 5.0-, 7.0- and 8.0-SOB contained extrinsic proteins (mainly 7S, 11S and γ-conglycinin) and oleosins (24 kDa, 18 kDa and 16 kDa), and pH 9.0-, 11.0-SOB only had oleosins. Extrinsic protein contents decreased from 83.9% at pH 5.0-SOB to 58.2% at pH 8.0-SOB, zeta potentials increased gradually from -21.7 mV to -14.67 mV and particle sizes decreased from 924 nm to 359 nm with increasing extraction pHs. In stomach digestion, oleosins of pH 5.0-, 7.0-, 8.0-SOB were hydrolyzed more rapidly than extrinsic proteins. All 24 kDa oleosins were hydrolyzed during 5 min, and 18 kDa oleosins were completely hydrolyzed at 30 min for all SOBs. Oleosins were hydrolyzed and produced <15 kDa pepsin-resistant peptides. In addition, some 7S and 11S of pH 5.0-, 7.0-, 8.0-SOB resisted pepsin hydrolysis. For all SOB emulsions, zeta potentials decreased and the droplets sizes increased significantly (P < 0.05) with the extension of gastric digestive time. The oil droplets of all SOB emulsions aggregated, and the coalescence of oil droplets more easily occurred in pH 9.0- and 11.0-SOB emulsions. During the intestinal phase, zeta potentials and diameters of the droplets decreased gradually, and the droplets of SOB emulsions dispersed according to the CLSM. FFA release rates of pH 5.0-, 7.0-, 8.0-SOB emulsions increased rapidly and then increased slowly, however, the release tendency of pH 9.0-, 11.0-SOB emulsions were opposite. Total FFA releases of pH 5.0-, 7.0-, 8.0-, 9.0-, 11.0-SOB emulsions were respectively 52.5%, 70.5%, 59%, 48.8%, 43.3% at 180 min. Therefore, the digestive behaviors of SOB emulsions extracted at different pHs were different. This study will provide guidance for SOB products.

Expanding the gene pool for soybean improvement with its wild relatives.

Genetic diversity is a cornerstone of crop improvement, However, cultivated soybean (Glycine max) has undergone several genetic bottlenecks, including domestication in China, the introduction of landraces to other areas of the world and, latterly, selective breeding, leading to low genetic diversity the poses a major obstacle to soybean improvement. By contrast, there remains a relatively high level of genetic diversity in soybean's wild relatives, especially the perennial soybeans (Glycine subgenus Glycine), which could serve as potential gene pools for improving soybean cultivars. Wild soybeans are phylogenetically diversified and adapted to various habitats, harboring resistance to various biotic and abiotic stresses. Advances in genome and transcriptome sequencing enable alleles associated with desirable traits that were lost during domestication of soybean to be discovered in wild soybean. The collection and conservation of soybean wild relatives and the dissection of their genomic features will accelerate soybean breeding and facilitate sustainable agriculture and food production.

Ancestral function but divergent epigenetic regulation of HAIKU2 reveals routes of seed developmental evolution.

Evolution is driven by various mechanisms. A directional increase in the embryo to endosperm ratio is an evolutionary trend within the angiosperms. The endosperm constitutes a major portion of the seed volume in Poales and some dicots. However, in other dicots such as Arabidopsis and soybean, the endosperm proliferates early, followed by embryo growth to replace the endosperm. The Arabidopsis leucine-rich repeat receptor protein kinase AtHAIKU2 (AtIKU2) is a key regulator of early endosperm proliferation. In this study, we found that IKU2s from Brachypodium, rice, and soybean can complement the abnormal seed developmental phenotype of Atiku2, while AtIKU2 also rescues the defective endosperm proliferation in the Brachypodium BdIKU2 knockout mutant seeds. AtIKU2 and soybean GmIKU2 are actively expressed a few days after fertilization. Thereafter, expression of AtIKU2 is suppressed by the FIS-PRC2 complex-mediated H3K27me3. The soybean GmIKU2 locus is also enriched with H3K27me3 marks. The histone methyltransferase AtMEA is unique to Brassicaceae, but one GmSWN in soybean plays a similar role in seed development as AtMEA. By contrast, the BdIKU2 and rice OsIKU2 loci are continuously expressed and are devoid of H3K27me3 marks. Taken together, these results suggest that IKU2 genes retain an ancestral function, but the duration of their expression that is controlled by PRC2-mediated epigenetic silencing contributes to silenced or persistent endosperm proliferation in different species. Our study reveals an epigenetic mechanism that drives the development of vastly different seed ontogenies.

Time-series transcriptome comparison reveals the gene regulation network under salt stress in soybean (Glycine max) roots.

BACKGROUND: Soil salinity is a primary factor limiting soybean (Glycine max) productivity. Breeding soybean for tolerance to high salt conditions is therefore critical for increasing yield. To explore the molecular mechanism of soybean responses to salt stress, we performed a comparative transcriptome time-series analysis of root samples collected from two soybean cultivars with contrasting salt sensitivity. RESULTS: The salt-tolerant cultivar 'Qi Huang No.34' (QH34) showed more differential expression of genes than the salt-sensitive cultivar 'Dong Nong No.50' (DN50). We identified 17,477 genes responsive to salt stress, of which 6644 exhibited distinct expression differences between the two soybean cultivars. We constructed the corresponding co-expression network and performed Gene Ontology term and Kyoto Encyclopedia of Genes and Genomes pathway enrichment analysis. The results suggested that phytohormone signaling, oxidoreduction, phenylpropanoid biosynthesis, the mitogen-activated protein kinase pathway and ribosome metabolism may play crucial roles in response to salt stress. CONCLUSIONS: Our comparative analysis offers a comprehensive understanding of the genes involved in responding to salt stress and maintaining cell homeostasis in soybean. The regulatory gene networks constructed here also provide valuable molecular resources for future functional studies and breeding of soybean with improved tolerance to salinity.

Phylogenomics of the genus Glycine sheds light on polyploid evolution and life-strategy transition.

Polyploidy and life-strategy transitions between annuality and perenniality often occur in flowering plants. However, the evolutionary propensities of polyploids and the genetic bases of such transitions remain elusive. We assembled chromosome-level genomes of representative perennial species across the genus Glycine including five diploids and a young allopolyploid, and constructed a Glycine super-pangenome framework by integrating 26 annual soybean genomes. These perennial diploids exhibit greater genome stability and possess fewer centromere repeats than the annuals. Biased subgenomic fractionation occurred in the allopolyploid, primarily by accumulation of small deletions in gene clusters through illegitimate recombination, which was associated with pre-existing local subgenomic differentiation. Two genes annotated to modulate vegetative-reproductive phase transition and lateral shoot outgrowth were postulated as candidates underlying the perenniality-annuality transition. Our study provides insights into polyploid genome evolution and lays a foundation for unleashing genetic potential from the perennial gene pool for soybean improvement.

Genome-Wide Identification, Characterization and Expression Analysis of Soybean CHYR Gene Family.

The CHYR (CHY ZINC-FINGER AND RING FINGER PROTEIN) proteins have been functionally characterized in iron regulation and stress response in Arabidopsis, rice and Populus. However, their roles in soybean have not yet been systematically investigated. Here, in this study, 16 GmCHYR genes with conserved Zinc_ribbon, CHY zinc finger and Ring finger domains were obtained and divided into three groups. Moreover, additional 2-3 hemerythrin domains could be found in the N terminus of Group III. Phylogenetic and homology analysis of CHYRs in green plants indicated that three groups might originate from different ancestors. Expectedly, GmCHYR genes shared similar conserved domains/motifs distribution within the same group. Gene expression analysis uncovered their special expression patterns in different soybean tissues/organs and under various abiotic stresses. Group I and II members were mainly involved in salt and alkaline stresses. The expression of Group III members was induced/repressed by dehydration, salt and alkaline stresses, indicating their diverse roles in response to abiotic stress. In conclusion, our work will benefit for further revealing the biological roles of GmCHYRs.

A transposon-mediated reciprocal translocation promotes environmental adaptation but compromises domesticability of wild soybeans.

Large structural variations frequently occur in higher plants; however, the impact of such variations on plant diversification, adaptation and domestication remains elusive. Here, we mapped and characterised a reciprocal chromosomal translocation in soybeans and assessed its effects on diversification and adaptation of wild (Glycine soja) and semiwild (Glycine gracilis) soybeans, and domestication of cultivated soybean (Glycine max), by tracing the distribution of the translocation in the USDA Soybean Germplasm Collection and population genetics analysis. We demonstrate that the translocation occurred through CACTA transposon-mediated chromosomal breakage in wild soybean c. 0.34 Ma and is responsible for semisterility in translocation heterozygotes and reduces their reproductive fitness. The translocation has differentiated Continental (i.e. China and Russia) populations from Maritime (i.e. Korea and Japan) populations of G. soja and predominately adapted to cold and dry climates. Further analysis revealed that the divergence of G. max from G. soja predates the translocation event and that G. gracilis is an evolutionary intermediate between G. soja and G. max. Our results highlight the effects of a chromosome rearrangement on the processes leading to plant divergence and adaptation, and provides evidence that suggests G. gracilis, rather than G. soja, as the ancestor of cultivated soybean.

Pedigree-based genetic dissection of quantitative loci for seed quality and yield characters in improved soybean.

As soybean plays an indispensable role in the supply of vegetable oil and protein, balancing the relationship between seed quality and yield traits according to human demand has become an important breeding goal for soybean improvement. Here, 256 intraspecific recombinant inbred lines (RILs), derived from a cross between Qi Huang No.34 (QH34) and Ji Dou No.17 (JD17), were used for quantitative trait loci (QTLs) mapping with remarkable four chemical and physical properties with a purpose for exploring the distribution of excellent alleles in germplasm resources in China. A total of 25 QTLs were detected, of which 10 QTLs inherited the alleles from the parent QH34. Pedigree research on favorable alleles on these QTLs showed the process of excellent alleles pyramided into QH34. Meta-analysis of the 25 QTLs by comparing with existed QTLs in previous study identified 17 novel QTLs. QTLs with pleiotropic effects have been detected. Furthermore, three representative elite recombinant inbred lines in different locations that have great potential in soybean breeding were selected, and finally, four seed weight-related candidate genes were identified. The discovery of these QTLs provides a new guidance for combining the diversity and rarity of germplasm resources, which can effectively increase population genetic diversity and broaden genetic basis of varieties. Supplementary Information: The online version contains supplementary material available at 10.1007/s11032-021-01211-6.

Genetic dissection of seed appearance quality using recombinant inbred lines in soybean.

Soybean seed appearance quality greatly affects the marketability. The objective of this study was to identify the quantitative trait loci (QTLs) that control the appearance quality of soybean seeds. A total of 256 recombinant inbred lines from Qi Huang No.34 × Ji Dou No.17 were utilized for QTL mapping. We innovatively applied a machine vision system to quantify the seed appearance of each line. As a result of QTL mapping, a total of 145 QTLs for the machine vision parameters were detected across three environments. We integrated QTLs mapped overlapped and obtained 16 QTL hotspots in total. Of these hotspots, hotspot-4-1 was suggested to be a major locus controlling seed size, and hotspot-15 was identified to affect the seed color and texture. The mapping for principal components of the seed appearance also supported it. This study comprehensively dug up the QTLs for seed appearance quality of soybean cultivars while providing an efficient method for phenotyping of seed appearance. These results would contribute to dissecting the genetic bases of seed appearance quality for the improvement of soybean. Supplementary Information: The online version contains supplementary material available at 10.1007/s11032-021-01262-9.

MPK14-mediated auxin signaling controls lateral root development via ERF13-regulated very-long-chain fatty acid biosynthesis.

Auxin plays a critical role in lateral root (LR) formation. The signaling module composed of auxin-response factors (ARFs) and lateral organ boundaries domain transcription factors mediates auxin signaling to control almost every stage of LR development. Here, we show that auxin-induced degradation of the APETALA2/Ethylene Responsive Factor (AP2/ERF) transcription factor ERF13, dependent on MITOGEN-ACTIVATED PROTEIN KINASE MPK14-mediated phosphorylation, plays an essential role in LR development. Overexpression of ERF13 results in restricted passage of the LR primordia through the endodermal layer, greatly reducing LR emergence, whereas the erf13 mutants showed an increase in emerged LR. ERF13 inhibits the expression of 3-ketoacyl-CoA synthase16 (KCS16), which encodes a fatty acid elongase involved in very-long-chain fatty acid (VLCFA) biosynthesis. Overexpression of KCS16 or exogenous VLCFA treatment rescues the LR emergence defects in ERF13 overexpression lines, indicating a role downstream of the auxin-MPK14-ERF13 signaling module. Collectively, our study uncovers a novel molecular mechanism by which MPK14-mediated auxin signaling modulates LR development via ERF13-regulated VLCFA biosynthesis.