Genetic Analysis of Soybean Flower Size Phenotypes Based on Computer Vision and Genome-Wide Association Studies.

The dimensions of organs such as flowers, leaves, and seeds are governed by processes of cellular proliferation and expansion. In soybeans, the dimensions of these organs exhibit a strong correlation with crop yield, quality, and other phenotypic traits. Nevertheless, there exists a scarcity of research concerning the regulatory genes influencing flower size, particularly within the soybean species. In this study, 309 samples of 3 soybean types (123 cultivar, 90 landrace, and 96 wild) were re-sequenced. The microscopic phenotype of soybean flower organs was photographed using a three-eye microscope, and the phenotypic data were extracted by means of computer vision. Pearson correlation analysis was employed to assess the relationship between petal and seed phenotypes, revealing a strong correlation between the sizes of these two organs. Through GWASs, SNP loci significantly associated with flower organ size were identified. Subsequently, haplotype analysis was conducted to screen for upstream and downstream genes of these loci, thereby identifying potential candidate genes. In total, 77 significant SNPs associated with vexil petals, 562 significant SNPs associated with wing petals, and 34 significant SNPs associated with keel petals were found. Candidate genes were screened by candidate sites, and haplotype analysis was performed on the candidate genes. Finally, the present investigation yielded 25 and 10 genes of notable significance through haplotype analysis in the vexil and wing regions, respectively. Notably, Glyma.07G234200, previously documented for its high expression across various plant organs, including flowers, pods, leaves, roots, and seeds, was among these identified genes. The research contributes novel insights to soybean breeding endeavors, particularly in the exploration of genes governing organ development, the selection of field materials, and the enhancement of crop yield. It played a role in the process of material selection during the growth period and further accelerated the process of soybean breeding material selection.

Morphological characterization and transcriptome analysis of rolled and narrow leaf mutant in soybean.

BACKGROUND: In plants, the leaf functions as a solar panel, where photosynthesis converts carbon dioxide and water into carbohydrates and oxygen. In soybean, leaf type traits, including leaf shape, leaf area, leaf width, and leaf width so on, are considered to be associated with yield. In this study, we performed morphological characterization, transcriptome analysis, and endogenous hormone analysis of a rolled and narrow leaf mutant line (rl) in soybean. RESULTS: Compared with wild type HX3, mutant line rl showed rolled and narrower leaflet, and smaller leaf, meanwhile rl also performed narrower pod and narrower seed. Anatomical analysis of leaflet demonstrated that cell area of upper epidermis was bigger than the cell area of lower epidermis in rl, which may lead rolled and narrow leaf. Transcriptome analysis revealed that several cytokinin oxidase/dehydrogenase (CKX) genes (Glyma.06G028900, Glyma.09G225400, Glyma.13G104700, Glyma.14G099000, and Glyma.17G054500) were up-regulation dramatically, which may cause lower cytokinin level in rl. Endogenous hormone analysis verified that cytokinin content of rl was lower. Hormone treatment results indicated that 6-BA rescued rolled leaf enough, rescued partly narrow leaf. And after 6-BA treatment, the cell area was similar between upper epidermis and lower epidermis in rl. Although IAA content and ABA content were reduced in rl, but exogenous IAA and ABA didn't affect leaf type of HX3 and rl. CONCLUSIONS: Our results suggest abnormal cytokinin metabolism caused rolled and narrow leaf in rl, and provide valuable clues for further understanding the mechanisms underlying leaf development in soybean.

Identification of Quantitative Trait Loci Controlling Root Morphological Traits in an Interspecific Soybean Population Using 2D Imagery Data.

Roots are the hidden and most important part of plants. They serve as stabilizers and channels for uptaking water and nutrients and play a crucial role in the growth and development of plants. Here, two-dimensional image data were used to identify quantitative trait loci (QTL) controlling root traits in an interspecific mapping population derived from a cross between wild soybean 'PI366121' and cultivar 'Williams 82'. A total of 2830 single-nucleotide polymorphisms were used for genotyping, constructing genetic linkage maps, and analyzing QTLs. Forty-two QTLs were identified on twelve chromosomes, twelve of which were identified as major QTLs, with a phenotypic variation range of 36.12% to 39.11% and a logarithm of odds value range of 12.01 to 17.35. Two significant QTL regions for the average diameter, root volume, and link average diameter root traits were detected on chromosomes 3 and 13, and both wild and cultivated soybeans contributed positive alleles. Six candidate genes, Glyma.03G027500 (transketolase/glycoaldehyde transferase), Glyma.03G014500 (dehydrogenases), Glyma.13G341500 (leucine-rich repeat receptor-like protein kinase), Glyma.13G341400 (AGC kinase family protein), Glyma.13G331900 (60S ribosomal protein), and Glyma.13G333100 (aquaporin transporter) showed higher expression in root tissues based on publicly available transcriptome data. These results will help breeders improve soybean genetic components and enhance soybean root morphological traits using desirable alleles from wild soybeans.

Tradeoffs among root functional traits for phosphorus acquisition in 13 soybean genotypes contrasting in mycorrhizal colonization.

BACKGROUND AND AIMS: Plants have adapted to acquire phosphorus (P) primarily through advantageous root morphologies, responsive physiological pathways and associations with mycorrhizal fungi. Yet, to date, little information exists on how variation in arbuscular mycorrhizal (AM) colonization is coordinated with root morphological and physiological traits to enhance P acquisition. METHODS: Thirteen root functional traits associated with P acquisition were characterized at full bloom stage in pot cultures under low soil P availability conditions for 13 soybean genotypes contrasting in AM colonization. KEY RESULTS: Significant variation in root functional traits was observed in response to low P stress among the 13 tested soybean genotypes contrasting in AM colonization. Genotypes with low AM colonization exhibited greater root proliferation but with less advantageous root physiological characteristics for P acquisition. In contrast, genotypes with high AM colonization exhibited less root growth but higher phosphatase activities and carboxylate content in the rhizosheath. Root dry weights, and contents of carbon and P were positively correlated with root morphological traits of different root orders and whole root systems, and were negatively correlated with AM colonization of fine roots and whole root systems, as well as rhizosheath phosphatase activities and carboxylate contents. These results taken in combination with a significant positive correlation between plant P content and root morphological traits indicate that root morphological traits play a primary role in soybean P acquisition. CONCLUSIONS: The results suggest that efficient P acquisition involves tradeoffs among carbon allocation to root proliferation, mycorrhizal symbiosis or P-mobilizing exudation. Complementarity and complexity in the selection of P acquisition strategies was notable among soybean genotypes contrasting in AM colonization, which is closely related to plant C budgeting.

Morphological and chemical plant traits associated with feeding non-preference to adult of Diabrotica speciosa (Coleoptera: Chrysomelidae) in soybean genotypes.

Diabrotica speciosa is an important pest of several crops in South America, including soybeans. Adults cause severe defoliation in soybean plants, and damage is significant when cotyledons are attacked. This study evaluated feeding non-preference to D. speciosa adults using 10 soybean genotypes, testing (i) 15-day-old whole plants and (ii) leaf disks of 60-day-old plants, through assessments of soybean attractiveness and leaf area consumed (LAC). Foliar contents of flavonoids and nutrients, and leaf trichome density were quantified for potential correlations with soybean resistance to adult of D. speciosa. In the whole young-plant experiment, under free-choice conditions, the lowest LAC was observed in IAC 100 and PI 227687. In no-choice, PI 227687 and IGRA RA 626 RR showed lower LAC than the other genotypes. In the leaf disk test, in free-choice, the genotypes IAC 100, PI 274454, PI 227687, DM 339, and BR 16 were the least preferred by adult of D. speciosa. In no-choice, PI 274454 was one of the least preferred, similarly to IGRA RA 626 RR, Dowling, and PI 227687. In the whole plant experiment, a high rutin content and low amounts of zinc, calcium, sulfur and manganese were associated with less consumption of D. speciosa on leaves of resistant genotypes. In contrast, in the leaf disk test there was a significant influence of trichomes in soybean resistance to the pest. In conclusion, the PI lines herein assessed are also promising sources for developing cultivars resistant to D. speciosa.

GmPIN1-mediated auxin asymmetry regulates leaf petiole angle and plant architecture in soybean.

Crop breeding during the Green Revolution resulted in high yields largely due to the creation of plants with semi-dwarf architectures that could tolerate high-density planting. Although semi-dwarf varieties have been developed in rice, wheat and maize, none was reported in soybean (Glycine max), and few genes controlling plant architecture have been characterized in soybean. Here, we demonstrate that the auxin efflux transporter PINFORMED1 (GmPIN1), which determines polar auxin transport, regulates the leaf petiole angle in soybean. CRISPR-Cas9-induced Gmpin1abc and Gmpin1bc multiple mutants displayed a compact architecture with a smaller petiole angle than wild-type plants. GmPIN1 transcripts and auxin were distributed asymmetrically in the petiole base, with high levels of GmPIN1a/c transcript and auxin in the lower cells, which resulted in asymmetric cell expansion. By contrast, the (iso)flavonoid content was greater in the upper petiole cells than in the lower cells. Our results suggest that (iso)flavonoids inhibit GmPIN1a/c expression to regulate the petiole angle. Overall, our study demonstrates that a signal cascade that integrates (iso)flavonoid biosynthesis, GmPIN1a/c expression, auxin accumulation, and cell expansion in an asymmetric manner creates a desirable petiole curvature in soybean. This study provides a genetic resource for improving soybean plant architecture.

Biophysical characteristics of soybean estimated by remote sensing associated with artificial intelligence

The biophysical characteristics of vegetative canopies, such as biomass, height, and canopy diameter, are of paramount importance for the study of the development and productive behavior of crops. Faced with a scarcity of studies aimed at estimating these parameters, the objective of this study was to evaluate the performance of artificial neural networks (ANNs) applied to Proximal Remote Sensing (PRS) to estimate biophysical characteristics of soybean culture. The data used to train and validate the ANNs came from an experiment composed of 65 plots with 30 x 30 m mesh, its development was carried out in the 2016/2017 crop in the Brazilian agricultural area. The evaluations were carried out at 30, 45, 60, and 75 days after sowing (DAS), monitoring the spatial and temporal variability of the biophysical characteristics of the soybean crop. Vegetation indexes were collected using canopy sensors. The accuracy and precision were determined by the coefficient of determination (R2) and the error of the forecasts by MAPE (Mean Absolute Percentage Error). PRS and ANNs showed high potential for application in agriculture, since they obtained good performance in the estimation of height (R2 = 0.89) and canopy diameter (R2 = 0.96), being fresh biomass (R2 =0.98) and dry biomass (R2 = 0.97) were the best-estimated variables.

GmWRKY46, a WRKY transcription factor, negatively regulates phosphorus tolerance primarily through modifying root morphology in soybean.

Phosphorus (P) deficiency affects soybean growth and development, resulting in significant reduction of yields. However, the regulatory mechanism of P deficiency tolerance in soybean is still largely unclear. WRKY transcription factors are a family of regulators involved in a variety of abiotic stresses in plants while rarely reported in P deficiency. Here, we demonstrated that a soybean GmWRKY46 gene, belonging to group III of WRKY TF family, was involved in the regulation of P deficiency tolerance in soybean. The expression of GmWRKY46 in low P sensitive soybean varieties was significantly higher than that in tolerant soybean varieties. It was primarily expressed in roots and strongly induced by P deprivation. GmWRKY46 was localized in the nucleus. Compared with the control expressing the empty vector, overexpression of GmWRKY46 in soybean hairy roots exhibited more sensitive phenotypes to low P stress, while the RNA interfered GmWRKY46 significantly enhanced P deficiency tolerance by increasing the proliferation, elongation and P absorption efficiency of hairy roots. Expression patterns of a number of P-responsive genes (GmPht1;1, GmPht1;4, GmPTF1, GmACP1, GmPAP21 and GmExpansin-A7) were altered in both overexpression and gene silenced plants. The results provided a novel insight into how soybean responds to low P stress and new gene that may be used to improve soybean low P tolerance through gene editing approach.

Genome-wide association studies of plant architecture-related traits and 100-seed weight in soybean landraces.

BACKGROUND: Plant architecture-related traits (e.g., plant height (PH), number of nodes on main stem (NN), branch number (BN) and stem diameter (DI)) and 100-seed weight (100-SW) are important agronomic traits and are closely related to soybean yield. However, the genetic basis and breeding potential of these important agronomic traits remain largely ambiguous in soybean (Glycine max (L.) Merr.). RESULTS: In this study, we collected 133 soybean landraces from China, phenotyped them in two years at two locations for the above five traits and conducted a genome-wide association study (GWAS) using 82,187 single nucleotide polymorphisms (SNPs). As a result, we found that a total of 59 SNPs were repeatedly detected in at least two environments. There were 12, 12, 4, 4 and 27 SNPs associated with PH, NN, BN, DI and 100-SW, respectively. Among these markers, seven SNPs (AX-90380587, AX-90406013, AX-90387160, AX-90317160, AX-90449770, AX-90460927 and AX-90520043) were large-effect markers for PH, NN, BN, DI and 100-SW, and 15 potential candidate genes were predicted to be in linkage disequilibrium (LD) decay distance or LD block. In addition, real-time quantitative PCR (qRT-PCR) analysis was performed on four 100-SW potential candidate genes, three of them showed significantly different expression levels between the extreme materials at the seed development stage. Therefore, Glyma.05 g127900, Glyma.05 g128000 and Glyma.05 g129000 were considered as candidate genes with 100-SW in soybean. CONCLUSIONS: These findings shed light on the genetic basis of plant architecture-related traits and 100-SW in soybean, and candidate genes could be used for further positional cloning.

Altered chromatin architecture and gene expression during polyploidization and domestication of soybean.

Polyploidy or whole-genome duplication (WGD) is widespread in plants and is a key driver of evolution and speciation, accompanied by rapid and dynamic changes in genomic structure and gene expression. The 3D structure of the genome is intricately linked to gene expression, but its role in transcription regulation following polyploidy and domestication remains unclear. Here, we generated high-resolution (∼2 kb) Hi-C maps for cultivated soybean (Glycine max), wild soybean (Glycine soja), and common bean (Phaseolus vulgaris). We found polyploidization in soybean may induce architecture changes of topologically associating domains and subsequent diploidization led to chromatin topology alteration around chromosome-rearrangement sites. Compared with single-copy and small-scale duplicated genes, WGD genes displayed more long-range chromosomal interactions and were coupled with higher levels of gene expression and chromatin accessibilities but void of DNA methylation. Interestingly, chromatin loop reorganization was involved in expression divergence of the genes during soybean domestication. Genes with chromatin loops were under stronger artificial selection than genes without loops. These findings provide insights into the roles of dynamic chromatin structures on gene expression during polyploidization, diploidization, and domestication of soybean.

Environmental and genetic regulation of plant height in soybean.

BACKGROUND: Shoot architecture is fundamentally crucial to crop growth and productivity. As a key component of shoot architecture, plant height is known to be controlled by both genetic and environmental factors, though specific details remain scarce. RESULTS: In this study, 308 representative soybean lines from a core collection and 168 F9 soybean progeny were planted at distinct field sites. The results demonstrated the presence of significant genotype × environment interaction (G × E) effects on traits associated with plant height in a natural soybean population. In total, 19 loci containing 51 QTLs (quantitative trait locus) for plant height were identified across four environments, with 23, 13 and 15 being QTLs for SH (shoot height), SNN (stem node number) and AIL (average internode length), respectively. Significant LOD ranging from 2.50 to 16.46 explained 2.80-26.10% of phenotypic variation. Intriguingly, only two loci, Loc11 and Loc19-1, containing 20 QTLs, were simultaneously detected across all environments. Results from Pearson correlation analysis and PCA (principal component analysis) revealed that each of the five agro-meteorological factors and four soil properties significantly affected soybean plant height traits, and that the corresponding QTLs had additive effects. Among significant environmental factors, AD (average day-length), AMaT (average maximum temperature), pH, and AN (available nitrogen) had the largest impacts on soybean plant height. Therefore, in spite of uncontrollable agro-meteorological factors, soybean shoot architecture might be remolded through combined efforts to produce superior soybean genetic materials while also optimizing soil properties. CONCLUSIONS: Overall, the comprehensive set of relationships outlined herein among environment factors, soybean genotypes and QTLs in effects on plant height opens new avenues to explore in work aiming to increase soybean yield through improvements in shoot architecture.

Anatomical changes in stem and root of soybean plants submitted to salt stress.

The soybean is a legume that is widely cultivated in many countries due to the high levels of protein and oil contained in its seed, and is used for human and animal nutrition. However, salinity affects more than 800 million hectares worldwide, limiting global agricultural production. The aim of this research was to evaluate the structural behaviour of the roots and stems under progressive salt stress, detailing the possible anatomical modifications to these organs in soybean plants during this stress. The plants were randomized into five treatments (0, 50, 100, 150 and 200 mm NaCl). All the root regions studied and exposed to 100 mm Na+ exhibited increases in the epidermis and endodermis and formation of lysogenic aerenchyma with increasing salinity, revealing the protective roles of these structures in reducing Na+ influx. In the stem, increases in the cortex and pith in the first internode subject to 100 mm Na+ suggest anatomical responses that aim to minimize oxidative stress. Soybean plants subjected to progressive salt stress (>50 mm Na+ ) avoided cavitation and loss of function linked to vessel elements, reducing the metaxylem in all the root and stem regions analysed. Finally, our results confirm anatomical changes to the roots and stems.

GmCRY1s modulate gibberellin metabolism to regulate soybean shade avoidance in response to reduced blue light.

Soybean is an important legume crop that displays the classic shade avoidance syndrome (SAS), including exaggerated stem elongation, which leads to lodging and yield reduction under density farming conditions. Here, we compared the effects of two shade signals, low red light to far-red light ratio (R:FR) and low blue light (LBL), on soybean status and revealed that LBL predominantly induces excessive stem elongation. We used CRISPR-Cas9-engineered Gmcry mutants to investigate the functions of seven cryptochromes (GmCRYs) in soybean and found that the four GmCRY1s overlap in mediating LBL-induced SAS. Light-activated GmCRY1s increase the abundance of the bZIP transcription factors STF1 and STF2, which directly upregulate the expression of genes encoding GA2 oxidases to deactivate GA1 and repress stem elongation. Notably, GmCRY1b overexpression lines displayed multiple agronomic advantages over the wild-type control under both dense planting and intercropping conditions. Our study demonstrates the integration of GmCRY1-mediated signals with the GA metabolic pathway in the regulation of LBL-induced SAS in soybean. It also provides a promising option for breeding lodging-resistant, high-yield soybean cultivars in the future.

Anatomical changes induced by isolates of Trichoderma spp. in soybean plants.

In the current work we evaluated the anatomical changes induced by T. harzianum and T. asperellum in two soybean cultivars, BRSGO Caiaponia and NA 5909 RG. Soybean production represents a growing market worldwide, and new methods aimed at increasing its productivity and yield are constantly being sought. Fungi of the genus Trichoderma have been widely used in agriculture as a promising alternative for the promotion of plant growth and for biological control of various pathogens. It is known that Trichoderma spp. colonize plant roots, but the anatomical changes that this fungus can cause are still less studied. Experiment was conducted in a greenhouse to collect leaves and soybean roots to perform analysis of growth parameters, enzymatic activity of defense-related enzymes and anatomical changes. It was observed that inoculation of Trichoderma spp. caused anatomical alterations, among them, increase in stomatal index at the abaxial leaf surface, thickness of the root cortex, thickness of adaxial epidermis, mean diameter of the vascular cylinder, thickness of the mesophyll, and thickness of the spongy parenchyma of the soybean plants. These results indicate that the alterations in these factors may be related to the process of plant resistance to pathogens, and better performance against adverse conditions. This study demonstrates that the anatomical study of plants is an important tool to show the effects that are induced by biological control agents.

High-throughput sequencing reveals the molecular mechanisms determining the stay-green characteristic in soybeans.

Senescence is an internally systematized degeneration process leading to death in plants. Leaf yellowing, one of the most prominent features of plant aging may lead to reduced crop yields. The molecular mechanism of responses to senescence in soybean leaves is not completely clear. In our research, two soybean varieties were selected with different stay-green traits: stay-green variety (BN106) and non-stay-green variety (KF14). RNA samples extracted from the leaves of two varieties were sequenced and compared using high-throughput sequencing. Six key enzyme genes in chlorophyll degradation pathways were studied to analyze the changes in their expression at seedling, flowering and maturation stage. Meanwhile, the construction of the genetic transformation process had been constructed to identify the function of putative gene by RNA-interference. A total of 4329 DEGs were involved in 52 functional groups and 254 KEGG pathways. Twelve genes encoding senescence-associated and inducible chloroplast stay-green protein showed significant differential expression. MDCase and PAO have a significant expression in BN106 that may be the key factors affecting the maintenance of green characteristics. In addition, the function of GmSGRs has been identified by genetic transformation. The loss of GmSGRs may cause soybean seeds to change from yellow to green. In summary, our results revealed fundamental information about the molecular mechanism of aging in soybeans with different stay-green characteristics. The work of genetic transformation lays a foundation for putative gene function studies that could contribute to postpone aging in soybeans.

Modification of soybean growth and abiotic stress tolerance by expression of truncated ERECTA protein from Arabidopsis thaliana.

ERECTA gene family encodes leucine-rich repeat receptor-like kinases that control major aspects of plant development such as elongation of aboveground organs, leaf initiation, development of flowers, and epidermis differentiation. To clarify the importance of ERECTA signaling for the development of soybean (Glycine max), we expressed the dominant-negative ERECTA gene from Arabidopsis thaliana that is truncated in the kinase domain (AtΔKinase). Expression of AtΔKinase in soybean resulted in the short stature, reduced number of leaves, reduced leaf surface area and enhanced branching in the transgenic plants. The transgenic AtΔKinase soybean plants exhibited increased tolerance to water deficit stress due to the reduction of total leaf area and reduced transpiration compared to the wild-type plants. Production of seeds in AtΔKinase lines was higher compared to wild type at regular conditions of cultivation and after exposure to drought stress. Transgenic seedlings expressing AtΔKinase were also able to withstand salt stress better than the wild-type. Established results demonstrated the significance of native soybean genes (GmER and GmERL) in development and stress response of soybean, and suggested that the truncated ERECTA gene of Arabidopsis thaliana can be used to manipulate the growth and stress response of different crop species.

LeafScope: A Portable High-Resolution Multispectral Imager for In Vivo Imaging Soybean Leaf.

Portable devices for measuring plant physiological features with their isolated measuring chamber are playing an increasingly important role in plant phenotyping. However, currently available commercial devices of this type, such as soil plant analysis development (SPAD) meter and spectrometer, are dot meters that only measure a small region of the leaf, which does not perfectly represent the highly varied leaf surface. This study developed a portable and high-resolution multispectral imager (named LeafScope) to in-vivo image a whole leaf of dicotyledon plants while blocking the ambient light. The hardware system is comprised of a monochrome camera, an imaging chamber, a lightbox with different bands of light-emitting diodes (LEDs) array, and a microcontroller. During measuring, the device presses the leaf to lay it flat in the imaging chamber and acquires multiple images while alternating the LED bands within seconds in a certain order. The results of an experiment with soybean plants clearly showed the effect of nitrogen and water treatments as well as the genotype differences by the color and morphological features from image processing. We conclude that the low cost and easy to use LeafScope can provide promising imaging quality for dicotyledon plants, so it has great potential to be used in plant phenotyping.

Toward a "Green Revolution" for Soybean.

Soybean (Glycine max), as an economically important food and oilseedcrop, is a major source of plant proteins and oils. Although considerable progress has been made in increasing the yields of rice, wheat, and maize through the "Green Revolution", little improvements have been made for soybean. With the increasing demand of soybean production and the rapid development of crop breeding technologies, time has come for this important crop to undergo a Green Revolution. Here, we briefly summarize the history of crop breeding and Green Revolution in other crops. We then discuss the possible directions and potential approaches toward achieving a Green Revolution for soybean. We provide our views and perspectives on how to breed new soybean varieties with improved yield.

High Throughput Phenotyping for Various Traits on Soybean Seeds Using Image Analysis.

Data phenotyping traits on soybean seeds such as shape and color has been obscure because it is difficult to define them clearly. Further, it takes too much time and effort to have sufficient number of samplings especially length and width. These difficulties prevented seed morphology to be incorporated into efficient breeding program. Here, we propose methods for an image acquisition, a data processing, and analysis for the morphology and color of soybean seeds by high-throughput method using images analysis. As results, quantitative values for colors and various types of morphological traits could be screened to create a standard for subsequent evaluation of the genotype. Phenotyping method in the current study could define the morphology and color of soybean seeds in highly accurate and reliable manner. Further, this method enables the measurement and analysis of large amounts of plant seed phenotype data in a short time, which was not possible before. Fast and precise phenotype data obtained here may facilitate Genome Wide Association Study for the gene function analysis as well as for development of the elite varieties having desirable seed traits.

Mapping developmental QTL for plant height in soybean [Glycine max (L.) Merr.] using a four-way recombinant inbred line population.

Plant height (PH) is an important trait in soybean, as taller plants may have higher yields but may also be at risk for lodging. Many genes act jointly to influence PH throughout development. To map the quantitative trait loci (QTL) controlling PH, we used the unconditional variable method (UVM) and conditional variable method (CVM) to analyze PH data for a four-way recombinant inbred line (FW-RIL) population derived from the cross of (Kenfeng14 × Kenfeng15) × (Heinong48 × Kenfeng19). We identified 7, 8, 16, 19, 15, 27, 17, 27, 22, and 24 QTL associated with PH at 10 developmental stages, respectively. These QTL mapped to 95 genomic regions. Among these QTL, 9 were detected using UVM and CVM, and 89 and 66 were only detected by UVM or CVM, respectively. In total, 36 QTL controlling PH were detected at multiple developmental stages and these made unequal contributions to genetic variation throughout development. Among 19 novel regions discovered in our study, 7 could explain over 10% of the phenotypic variation and contained only one single QTL. The unconditional and conditional QTL detected here could be used in molecular design breeding across the whole developmental procedure.