Polymorphism analysis of the chloroplast and mitochondrial genomes in soybean.

BACKGROUND: Soybean is an important protein- and oil-rich crop throughout the world. Much attention has been paid to its nuclear genome, which is bi-parentally inherited and associated with many important agronomical traits. However, less is known about the genomes of the semi-autonomous and essential organelles, chloroplasts and mitochondria, of soybean. RESULTS: Here, through analyzing the polymorphisms of these organelles in 2580 soybean accessions including 107 wild soybeans, we found that the chloroplast genome is more variable than the mitochondrial genome in terms of variant density. Consistent with this, more haplotypes were found in the chloroplast genome (44 haplotypes) than the mitochondrial genome (30 haplotypes). These haplotypes were distributed extremely unevenly with the top two haplotypes (CT1 and CT2 for chloroplasts, MT1 and MT2 for mitochondria) accounting for nearly 70 and 18% of cultivated soybean accessions. Wild soybeans also exhibited more diversity in organelle genomes, harboring 32 chloroplast haplotypes and 19 mitochondrial haplotypes. However, only a small percentage of cultivated soybeans shared cytoplasm with wild soybeans. In particular, the two most frequent types of cytoplasm (CT1/MT1, CT2/MT2) were missing in wild soybeans, indicating that wild soybean cytoplasm has been poorly exploited during breeding. Consistent with the hypothesis that soybean originated in China, we found that China harbors the highest cytoplasmic diversity in the world. The geographical distributions of CT1-CT3 and MT1-MT3 in Northeast China were not significantly different from those in Middle and South China. Two mitochondrial polymorphism sites, p.457333 (T > C) and p.457550 (G > A), were found to be heterozygous in most soybeans, and heterozygosity appeared to be associated with the domestication of cultivated soybeans from wild soybeans, the improvement of landraces to generate elite cultivated soybeans, and the geographic adaptation of soybean. CONCLUSIONS: The haplotypes of thousands of soybean cultivars should be helpful in evaluating the impact of cytoplasm on soybean performance and in breeding cultivars with the desired cytoplasm. Mitochondrial heterozygosity might be related to soybean adaptation, and this hypothesis needs to be further investigated.

Transcriptomal dissection of soybean circadian rhythmicity in two geographically, phenotypically and genetically distinct cultivars.

BACKGROUND: In soybean, some circadian clock genes have been identified as loci for maturity traits. However, the effects of these genes on soybean circadian rhythmicity and their impacts on maturity are unclear. RESULTS: We used two geographically, phenotypically and genetically distinct cultivars, conventional juvenile Zhonghuang 24 (with functional J/GmELF3a, a homolog of the circadian clock indispensable component EARLY FLOWERING 3) and long juvenile Huaxia 3 (with dysfunctional j/Gmelf3a) to dissect the soybean circadian clock with time-series transcriptomal RNA-Seq analysis of unifoliate leaves on a day scale. The results showed that several known circadian clock components, including RVE1, GI, LUX and TOC1, phase differently in soybean than in Arabidopsis, demonstrating that the soybean circadian clock is obviously different from the canonical model in Arabidopsis. In contrast to the observation that ELF3 dysfunction results in clock arrhythmia in Arabidopsis, the circadian clock is conserved in soybean regardless of the functional status of J/GmELF3a. Soybean exhibits a circadian rhythmicity in both gene expression and alternative splicing. Genes can be grouped into six clusters, C1-C6, with different expression profiles. Many more genes are grouped into the night clusters (C4-C6) than in the day cluster (C2), showing that night is essential for gene expression and regulation. Moreover, soybean chromosomes are activated with a circadian rhythmicity, indicating that high-order chromosome structure might impact circadian rhythmicity. Interestingly, night time points were clustered in one group, while day time points were separated into two groups, morning and afternoon, demonstrating that morning and afternoon are representative of different environments for soybean growth and development. However, no genes were consistently differentially expressed over different time-points, indicating that it is necessary to perform a circadian rhythmicity analysis to more thoroughly dissect the function of a gene. Moreover, the analysis of the circadian rhythmicity of the GmFT family showed that GmELF3a might phase- and amplitude-modulate the GmFT family to regulate the juvenility and maturity traits of soybean. CONCLUSIONS: These results and the resultant RNA-seq data should be helpful in understanding the soybean circadian clock and elucidating the connection between the circadian clock and soybean maturity.

GmFULa improves soybean yield by enhancing carbon assimilation without altering flowering time or maturity.

KEY MESSAGE: GmFULa improved soybean yield by enhancing carbon assimilation. Meanwhile, different from known yield-related genes, it did not alter flowering time or maturity. Soybean (Glycine max (L.) Merr.) is highly demanded by a continuously growing human population. However, increasing soybean yield is a major challenge. FRUITFULL (FUL), a MADS-box transcription factor, plays important roles in multiple developmental processes, especially fruit and pod development, which are crucial for soybean yield formation. However, the functions of its homologs in soybean are not clear. Here, through haplotype analysis, we found that one haplotype of the soybean homolog GmFULa (GmFULa-H02) is dominant in cultivated soybeans, suggesting that GmFULa-H02 was highly selected during domestication and varietal improvement of soybean. Interestingly, transgenic overexpression of GmFULa enhanced vegetative growth with more biomass accumulated and ultimately increased the yield but without affecting the plant height or changing the flowering time and maturity, indicating that it enhances the efficiency of dry matter accumulation. It also promoted the yield factors like branch number, pod number and 100-seed weight, which ultimately increased the yield. It increased the palisade tissue cell number and the chlorophyll content to promote photosynthesis and increase the soluble sugar content in leaves and fresh seeds. Furthermore, GmFULa were found to be sublocalized in the nucleus and positively regulate sucrose synthases (SUSs) and sucrose transporters (SUTs) by binding with the conserved CArG boxes in their promoters. Overall, these results showed GmFULa promotes the capacity of assimilation and the transport of the resultant assimilates to increase yield, and provided insights into the link between GmFULa and sucrose synthesis with transport-related molecular pathways that control seed yield.

GmFULa, a FRUITFULL homolog, functions in the flowering and maturation of soybean.

KEY MESSAGE: A FRUITFULL homolog GmFULa was cloned and found to play roles in the flowering and maturation of soybean. Soybean varieties exhibit great diversity in terms of flowering and maturation due to differences in their photoperiodic responses. The underlying mechanism remains unclear despite the fact that some upstream flowering genes have been studied. FRUITFULL (FUL) genes are one group of downstream flowering genes known to have major roles in reproductive transition, floral meristem identity, and floral organ identity. However, FUL homologs and their functions are poorly understood in soybean. Here, a soybean FUL homolog was cloned from the late-maturing photoperiod-sensitive soybean variety Zigongdongdou (ZGDD) and designated GmFULa. In ZGDD, GmFULa exhibited a terminal-preferential expression pattern, with higher expression in the root and shoot apices than in the middle parts. Diurnal rhythm analysis revealed that photoperiod regulates the GmFULa expression level but does not alter its diurnal rhythm. ZGDD was maintained under different photoperiod conditions (long day, LD; short day, SD; LD after 13 short days, SD13-LD) to assess GmFULa expression in newly expanded leaves and in the shoot apex. From this analysis, GmFULa expression was detected in the floral meristem, floral organs and their primordia; trifoliate leaves; and the inflorescence meristem, with the expression levels induced by SD and inhibited by LD. GmFULa expression was also associated with maturity in seven soybean varieties with different photoperiod sensitivities. Therefore, photoperiod conditions affect the expression level of GmFULa but not its diurnal rhythm. The gene plays pleiotropic roles in reproductive transition, flowering, and leaf development and is associated with maturity in soybean.

Allelic combinations of soybean maturity Loci E1, E2, E3 and E4 result in diversity of maturity and adaptation to different latitudes.

Soybean cultivars are extremely diverse in time to flowering and maturation as a result of various photoperiod sensitivities. The underlying molecular genetic mechanism is not fully clear, however, four maturity loci E1, E2, E3 and E4 have been molecularly identified. In this report, cultivars were selected with various photoperiod sensitivities from different ecological zones, which covered almost all maturity groups (MG) from MG 000 to MG VIII and MG X adapted from latitude N 18° to N 53°. They were planted in the field under natural daylength condition (ND) in Beijing, China or in pots under different photoperiod treatments. Maturity-related traits were then investigated. The four E maturity loci were genotyped at the molecular level. Our results suggested that these four E genes have different impacts on maturity and their allelic variations and combinations determine the diversification of soybean maturity and adaptation to different latitudes. The genetic mechanisms underlying photoperiod sensitivity and adaptation in wild soybean seemed unique from those in cultivated soybean. The allelic combinations and functional molecular markers for the four E loci will significantly assist molecular breeding towards high productivity.

GmFT2a polymorphism and maturity diversity in soybeans.

BACKGROUND: Soybean is a short-day crop of agricultural, ecological, and economic importance. The sensitive photoperiod responses significantly limit its breeding and adaptation. GmFT2a, a putative florigen gene with different transcription profiles in two cultivars (late-maturing Zigongdongdou and early-maturing Heihe 27) with different maturity profiles, is key to flowering and maturation. However, up to now, its role in the diverse patterns of maturation in soybeans has been poorly understood. METHODS: Eighty varieties, including 19 wild accessions, covering 11 of all 13 maturity groups, were collected. They were planted in pots and maintained under different photoperiodicity conditions (SD, short day; LD, long day; and ND, natural day). The day to first flowering was recorded and the sensitivity to photoperiod was investigated. Polymorphisms in the GmFT2a coding sequence were explored by searching the known SNP database (NCBI dbSNP). The GmFT2a promoter regions were then cloned from these varieties and sequenced. Further polymorphism and association analyses were conducted. RESULTS: These varieties varied greatly in time to first flowering under ND and exhibited a consecutive distribution of photoperiod sensitivity, which suggested that there is rich diversity in flowering time. Furthermore, although GmFT2a had only one known synonymous SNP in the coding sequence, there were 17 haplotypes of the GmFT2a promoter region, HT06 of which was extremely abundant. Further association analysis found some SNPs that might be associated with day to first flowering and photoperiod sensitivity. CONCLUSION: Although GmFT2a is a key flowering gene, GmFT2a polymorphism does not appear to be responsible for maturity diversity in soybean.