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    Topic review

    Korean Wild Soybeans

    Subjects: Plant Sciences
    View times: 10
    Submitted by: Muhammad Amjad Nawaz

    Definition

    Domesticated crops suffer from major genetic bottlenecks while wild relatives retain higher genomic diversity. Wild soybean (Glycine soja Sieb. & Zucc.) is the presumed ancestor of cultivated soybean (Glycine max [L.] Merr.), and is an important genetic resource for soybean improvement. Among the East Asian habitats of wild soybean (China, Japan, Korea, and Northeastern Russia), the Korean peninsula is of great importance based on archaeological records, domestication history, and higher diversity of wild soybeans in the region. The collection and conservation of these wild soybean germplasms should be put on high priority. Chung’s Wild Legume Germplasm Collection maintains more than 10,000 legume accessions with an intensive and prioritized wild soybean germplasm collection (>6000 accessions) guided by the international code of conduct for plant germplasm collection and transfer. The center holds a library of unique wild soybean germplasms collected from East Asian wild habitats including the Korean mainland and nearby islands. The collection has revealed interesting and useful morphological, biochemical, and genetic diversity. This resource could be utilized efficiently in ongoing soybean improvement programs across the globe.

    1. Introduction

    Soybean (Glycine max [L.] Merr.) is an economically important legume worldwide with diverse applications in food, feed, and biofuel industries [1][2]. Increasing per capita food consumption and limited land and water resources, coupled with a growing population and changing dietary habits, are projected to escalate the pressure on global food supplies [3][4]. It is estimated that food consumption is growing faster than production, particularly in Asia, which will lead to grain and soybean deficits. This deficit in food supply must be addressed through a global effort. Additionally, global climate change, together with the above-mentioned factors, has outpaced the forecasted crop yields, while at the same time having a negative effect on the state of diversity in the farmer’s field [5]. To alleviate negative effects due to climate change on crops and ecosystems and to increase crop yields, the resilience and sustainability of current agricultural systems have become pressing issues [6]. The development of new soybean varieties and cropping systems with the ability to adapt to a changing environment and new socio-economic conditions is crucial for yield enhancement. While the genetic improvement of crops plays an important role, the genetic base of major crops is narrow, due to genetic bottlenecks and human selection [7]. Therefore, it is essential to investigate novel sources of genetic diversity and to expand gene pools by exploiting crop wild relatives (CWRs) [8][9][10]. The search for novel genetic diversity will result in an increased demand for novel breeding material from the world’s gene banks.
    The wild (Glycine soja Sieb. & Zucc.) and cultivated (G. max) soybeans are both annual plants belonging to one of the two subgenera of the genus Glycine and are distinguished from other wild perennial Glycine species. It has been shown that wild soybean can be used in the genetic improvement of cultivated soybean [11].
    The taxonomic and phylogenetic knowledge based on the concept of gene pools provides important information for wild x cultivated soybean hybridization. The primary gene pool (GP-1) includes G. max and its wild progenitor, G. soja, resulting in the production of vigorous hybrids, normal meiotic chromosome pairing, and fertile offspring. As far as secondary (GP-2) and tertiary (GP-3) gene pools of G. max are concerned, all the 26 perennial species are included in GP-3, while no Glycine species has been found in GP-2 [11]. Some researchers have previously attempted to hybridize G. max with five GP-3 member species, but their success was limited [12]. Among GP-3 members, only three perennial species (Glycine argyrea, Glycine canescens, and Glycine tomentella) have been successfully hybridized with cultivated soybean. The F1 hybrids were either sterile or could not develop beyond the amphidiploid stage [13]. Further information could be found in the paper by Singh ([14] and references therein).
    Wild soybean accessions, the wild relatives of cultivated soybean, are mainly found in East Asia [15]. They harbor abundant and unique gene/allele resources, due to their widespread distributions in complex geographic topographies with diverse microclimates [16]. Cultivated soybeans have lower adaptive potential to climatic changes and reduced genetic diversity when compared to wild soybeans [3][17]. More than half of the genetic variations in soybean were lost due to habitat fragmentation and genetic bottlenecks [18]. Therefore, wild soybeans are important genetic resources for gaining a deeper understanding of the genetic diversity, genomic variations, and environmental adaptations of soybean [10][19].
    Preliminary assessments of germplasm conservation in many parts of the world have revealed substantial gaps, which should be filled by expanding the collection of wild soybean germplasm resources from the main distribution areas [20][21]. Such efforts are being adopted by many countries and germplasm centers around the globe, including Korea, which is one of the main distribution areas of wild soybeans.

    2. Distribution and Conservation of Wild Soybean in Korea

    2.1. Geographical Distribution of Wild Soybean in Korea

    Wild soybean grows in moist habitats from 0 to 2650 MASL (meters above sea level), spanning subtropical and subarctic zones (between 24° N and 53° N latitude) [16], and is restricted to East Asia. Occasionally, it can also be found in dry and salt-affected areas. In the Korean peninsula, wild soybeans are distributed throughout the mainland as well as on nearby islands. They can be found growing almost everywhere in Korea, including farmlands, roadsides, and river banks, and from mountain tops to deep valley bottoms. There are over 3000 islands off the coast of the Korean peninsula and nearly 400 are inhabited. A few of these inhabited islands have already been surveyed for wild soybean distribution [22].

    2.2. Archaeological Records and Background

    Despite the importance of soybean to the world economy, the history of soybean has been mostly restricted to phylogenetics and historical documents mentioning the domestication of soybean in East Asia. G. soja is the only wild member of the subgenus Soja native to the East Asian countries mentioned above [15][23]. Many reports suggest that soybean was first domesticated in China around 5000 years ago, but this statement is still controversial [24]. However, it is generally agreed that the ancient geographical region comprising China, Japan, and Korea (the CJK region) harbored one of the three pioneering societies to domesticate and cultivate this important legume [25]. In an attempt to rectify the domestication history of soybean, Lee et al. [26] pointed out that the differences between domesticated plants and their wild relatives/progenitors are not always clear, and therefore it is sometimes impossible to discern between wild and domesticated soybean in archaeological specimens which are not always well preserved. Despite their different appearances and growing conditions, cultivated soybean plants can cross-breed with wild soybean plants and produce fertile F1 hybrids, making it quite challenging to understand and interpret the archaeological seed records [11]. There is still an ongoing debate over the archaeological records in southeastern Korea and China ([26] and references therein).
    Figure 1 shows the archaeological sites in Korea. The Nam River valley contains multiple sites, e.g., Oun 1, Okbang 1, 2, 4, 6, 9, Sangchon B, and Pyeonggeodong, dating back to the Chulmun (8000~3500 CAL. B.P.), Mumun (~3500-2000 CAL. B.P.), and Three Kingdom Periods (2000~1200 CAL. B.P.). Other sites include Daundong in Ulsan and Dongsamdong shell midden in Busan. A detailed examination and comparison of archaeological seed records showed that the morphology of the seeds in the Korean soybean archaeological records resembles that of smaller domesticated varieties in more modern times [26][27] because there is a range of seed morphologies among soybean cultivars with ovate seeds. However, the report is based on archaeobotanical seed size measurements. A genomic approach could be more rewarding in terms of species identification. Another report [27] did not mention wild soybean in Korean flotation samples at all. Due to a higher number of landraces, as well as the presence of wild soybean in the area, the authors were not able to link any archaeological findings to a particular species (most probably G. max) or a soybean landrace. However, these studies confirm that the Korean peninsula is a region of great importance for early soybean domestication.
    Figure 1. Map showing the archaeological sites of soybeans in Korea. Bullets under each city show the names of the archaeological sites while the dates in brackets show the carbon dating information on charred soybean seeds. If there is no carbon dating of the soybean samples, then the reported chronological year is given [26][28].

    3. Collection of Wild Soybean Germplasm in Korea

    The second report on the state of the world’s plant genetic resources for food and agriculture stated that public awareness of the importance of crop diversity and the use of CWRs is growing in both developing and developed countries. A 20% increase in the number of global accessions conserved ex-situ has been observed since 1996. Major ex-situ soybean collections are reported to be present in China, Russia, Ukraine, and the United States of America, with a total of 229,944 accessions. However, 23% of the total collection is held by only two organizations, i.e., the Institute of Crop Germplasm Resources at The Chinese Academy of Agricultural Sciences (ICGR-CAAS) and SOY (USA) [5]. Among these soybean germplasm collections, the majority of the wild accessions are held by organizations in China, the USA, Korea, Japan, and Russia (Table 1). According to the second report on plant genetic resources, there has been a significant increase in nationally designated protected areas since 1928. However, the report did not include priority genetic reserve locations for wild soybean relatives. The most likely reason for the omission is that this report based its information on another report by Maxted and Kell which focused only on 12 major food crops, excluding soybean [29]. A recent report on global conservation priorities classified soybean wild relatives as medium priority [21]. This classification is based on the consideration given to the situations of all food crops. Therefore, a classification system based solely on the rankings within legumes or oilseeds could help advocate for a higher priority for conserving wild soybean.
    Table 1. Global soybean germplasm collections by the institute.
    Instcode Institute No. of Accessions % WS LR BL AC OT
    CHN001 ICGR-CAAS 32,021 14 21       79
    USA033 SOY 21,075 9 10 80 5 4 1
    KOR011 RDAGB-GRD 17,644 8 <1 45 5 1 50
    TWN001 AVRDC 15,314 7   <1   <1 100
    BRA014 CNPSO 11,800 5         100
    JPN003 NIAS 11,473 5 5 33 21   40
    RUS001 VIR 6439 3   9 40 41 11
    IND016 AICRP-Soybean 4022 2 <1       100
    CIV005 IDESSA 3727 2         100
    TWN006 TARI 2745 1         100
    DEU146 IPK 2661 1 1 23 53 23  
    ZWE003 CBICAU 2236 1         100
    IDN182 ICRR 2198 1 <1       100
    AUS048 ATCFC 2121 1 3 <1 38 52 6
    NGA039 IITA 1909 1   5 4 1 90
    FRA060 AMFO 1582 1         100
    THA005 FCRI-DA/TH 1510 1     100    
    MEX001 INIA-Iguala 1500 1         100
    PHL130 IPB-UPLB 1381 1   100      
    UKR001 IR 1288 1 3 1 21 72 3
    COL017 ICA/REGION 1 1235 1   <1 64 13 22
    SRB002 IFVCNS 1200 1       100  
    ROM002 ICCPT Fundul 1024 <1     62 38 <1
      Others (166) 81,839 36   11 4 27 51
      Total 229,944 100 6 17 7 13 56
    ICGR-CAAS Institute of Crop Germplasm Resources, Chinese Academy of Agricultural Sciences
    SOY Soybean Germplasm Collection, United States Department of Agriculture, Agricultural Research Services
    RDAGB-GRD Genetic Resources Division, National Institute of Agricultural Biotechnology, Rural Development Administration (Korea)
    AVRDC World Vegetable Centre (former Asian Vegetable Research and Development Centre)
    CNPSO Embrapa Soja (Brazil)
    NIAS National Institute of Agrobiological Sciences (Japan)
    VIR N.I. Vavilov All-Russian Scientific Research Institute of Plant Industry (Russian Federation)
    AICRP-Soybean All India Coordinated Research Project on Soybean (India)
    IDESSA Institut des Savanes (Côte d’Ivoire)
    TARI Taiwan Agricultural Research Institute
    IPK Genebank, Leibniz Institute of Plant Genetics and Crop Plant Research (Germany)
    CBICAU Crop Breeding Institute (Zimbabwe)
    ICRR Indonesian Centre for Rice Research
    ATCFC Australian Tropical Crops & Forages Genetic Resources Centre
    IITA International Institute of Tropical Agriculture
    AMFO G.I.E. Amelioration Fourragère (France)
    FCRI-DA/TH Field Crops Research Institute–Department of Agriculture (Thailand)
    INIA-Iguala Estación de Iguala, Instituto Nacional de Investigaciones Agrícolas (Mexico)
    IPB-UPLB Institute of Plant Breeding, College of Agriculture, University of the Philippines, Los Baños College (Philippines)
    IR Institute of Plant Production n.a. V.Y. Yurjev of UAAS (Ukraine)
    ICA/REGION 1 Corporación Colombiana de Investigación Agropecuaria Tibaitata (Colombia)
    IFVCNS Institute for Field and Vegetable Crops (Serbia)
    ICCPT Fundul Research Institute for Cereals and Technical Plants Fundulea (Romania)
    WS = Wild Relatives, LR = Landrace, BL = Breeding Line, AC = Advanced Cultivar, and OT = Others. Source: [5].
    Wild soybean, being able to hybridize with cultivated soybean, becomes an extended source of diversity to broaden the gene pool [11]. The utilization of wild soybean is expected to increase as a result of ongoing improvements in the information on the species’ genome and genetic diversity, and advances in breeding tools [21]. This expectation is based on the assumption that the wild accessions will be readily available for research and soybean breeding, which requires their conservation as germplasms in gene banks. Urbanization, road construction, deforestation, intensive agriculture, climate change, and soil erosion are all threatening wild soybean in its natural habitats. Since wild soybean is mainly distributed in the CJK region, greater conservation efforts should be made by the local authorities in charge of biological conservation in these countries. In East Asia, major wild soybean germplasm centers in these three countries include the Chinese Crop Germplasm Information System, in China, the National Institute of Agrobiological Sciences Genebank and the Legume Base, in Japan, and the National Agrobiodiversity Center and the Chung’s Wild Legume Germplasm Collection (CWLGC) (described in this work), in Korea. In Korea, there are >6000 accessions in CWLGC and 3229 accessions at the National Agrobiodiversity Center.
    Soybean breeding in Korea started in the early 1900s and, since then, more than 178 varieties/cultivars have been registered. Major soybean breeding work has been completed in the past three decades, particularly after the establishment of the National Agrobiodiversity Center in 1987 [30]. Cultivated soybean has received greater attention regarding germplasm collection when compared to wild soybean. This is quite logical because most of the germplasm centers were busy collecting, breeding, and documenting major crops during their early establishment. However, the recent trend of utilizing CWRs in breeding and preliminary assessments of the comprehensiveness of the conservation of CWRs in gene banks have reported substantial gaps [21]. A report by FAO on the state of diversity indicated that the number of accessions of CWRs has substantially increased since 1996. However, they are still under-represented [5]. Considering this scenario, as well as the limited number of wild soybean accessions collected from Korea, large-scale individual as well as collective efforts are needed to enhance wild soybean collections. Although the information is available about the conservation status of this species in Korea, the lack of information and access to wild soybean in North Korea is a big limitation. Table 2 details the conservation efforts by the CWLGC in Korea, placing its major focus on wild soybean collection.
    Table 2. List of wild legume germplasm accessions at Chung’s Wild Legume Germplasm Collection.
    Species No. of Accessions
    Glycine max (L.) Merr 600
    Glycine soja 6232
    Amphicarpaea edgeworthii 1300
    Vigna vexillata 810
    Rhynchosia volubilis 225
    Phaseolus nipponensis 700
    Other wild legumes
    Dunbaria villosa (Thunb.) Makino
    Vigna umbellate (Thumb.) Ohwi & Ohashi
    Vigna angularis var. nipponensis (ohwi) ohwi & ohashi
    Vigna angularis (Willd.) Ohwi & Ohashi
    700
    Total 10,567

    3.1. Chung’s Wild Legume Germplasm Collection

    Founded by Professor Gyuhwa Chung in 1983 and located in the Yeosu campus of the Chonnam National University, the CWLGC now possesses the most comprehensive collection of wild soybean in Korea. Guided by the international code of conduct for plant germplasm collecting and transfer (http://www.fao.org/3/x5586e/x5586e0k.htm), the main focus of this center is on the direct collection, acquisition, conservation, evaluation, characterization, documentation, and distribution of wild legume germplasms. CWLGC followed the guidelines of the National Agrobiodiversity Center, Genebank of Rural Development Administration, Korea, and became a local sub-bank in 2007. The list of species and the respective number of accessions available at CWLGC is shown in Table 2. The wild legume accessions were directly collected by the CWLGC team from Korea, Japan, China, and Russia, or acquired from Australia, India, Taiwan, and Zambia. The wild soybean collections include 5050 accessions originating from Korea.

    3.2. In situ Conservation of G. soja at CWLGC

    In the last 30 years, a major effort at CWLGC has been dedicated to wild soybean germplasm collection, acquisition, development, and characterization. In situ wild soybean germplasm propagation and characterization at CWLGC is done throughout the year and is ongoing. About 500-1000 accessions/year are propagated and characterized at two outdoor planting and propagation sites: Yeosu (Chonnam Province) and Jinju (Gyeongsangnam Province) (Figure 2).
    Figure 2. Wild soybean germplasm. (a) Wild soybean germplasm distribution and collection. (b) Different stages of in situ propagation and characterization at CWLGC.
    One important feature of CWLGC is that it holds wild soybean accessions collected from ~130 islands along the coast of the Korean peninsula. These accessions could be used to address questions related to the adaptations to microclimatic conditions in geographically isolated areas. With increasing land exploitations by humans and decreasing wild soybean habitats in Korea, CWLGC is focusing on extensive surveys and germplasm collections. The germplasm characterization performed at CWLGC has revealed interesting and useful morphological and biochemical variations. Examples of the morphological diversity in leaf shape and root architecture are shown in Figure 3.
    Figure 3. Morphological diversity of soybeans in the CWLGC collection. (a) Leaf shape variations in G. soja accessions. (b) Root structure variations in G. soja accessions. (c) Samples of the legume seed collection at CWLGC.
    The germplasm collection at CWLGC exhibits high genetic and morphological variations [22][26]. For instance, microsatellite marker-based genetic diversity has been investigated in the wild soybean at CWLGC that originated from the southern islands of Korea. Five hundred and thirty wild soybean accessions originating from China, Japan, and Korea included in the CWLGC have also been utilized for the successful identification of GmSALT3 haplotypes and the development of molecular markers related to salt tolerance [31].
    Moreover, the CWLGC germplasms were studied for their natural variations in saponin contents and were used to help identify different group-A acetylsaponin-deficient mutants [32][33][34][35]. The wild soybean soyasaponin mutants from CWLGC have been used to identify and characterize the genes involved in the saponin biosynthesis pathway, e.g., Sg-1 locus (CWS2133), sg-5 locus (CWS5095), and Sg-6 locus [36][34][37][38][39]. In addition to saponins, wild soybean accessions at the CWLGC have also been screened for the presence of Kunitz trypsin inhibitor polymorphism, alpha-linolenic acid concentration, and soyisoflavone profile diversity [40][41][42][43]. Furthermore, the germplasms have been used to functionally characterize two seed-specific flavonoid glycosyltransferases and a β-amyrin synthase gene [39][44].
    Another interesting discovery using the CWLGC collection is the identification of the W1 locus and the analysis of four new alleles at this locus associated with flower color in soybean [45]. A complete range of flower color has been observed in the CWLGC wild soybean collection and different flower color variants have been studied to help understand the genetic and molecular basis of flower color e.g., the pinkish-white flowers of accession CW13133 [46] (Figure 4).
    Figure 4. Demonstrations of the diversity in the CWLGC. (a) Flower color variations [45]. (b) Using Kunitz trypsin inhibitor, different KTi forms were detected in Korean G. soja accessions by non-denaturing PAGE. Lanes 1–9 are: Tia protein standard (Sigma), Tia, Tib, Tia/Tib, Tibi7-1, Tia, Tibi5, Tib, and Tia, respectively [40]. (c) Comparison of saponin composition phenotypes in seed hypocotyls in the Korean G. soja collection by thin-layer chromatography (TLC). Lanes 1–7 are the common phenotypes: Aa, Ab, AaBc, AbBc, Aa+a, AaBc+a, and AbBc+a types, respectively. Lanes 8, 9, and 10 are mutant phenotypes: AuAeBc (CWS0115), A0Bc+ag (CWS2133), and A0Bc-S (CWS5095) types, respectively [32].
    Being a major source of plant oils and proteins, the demand for soybean is escalating. Soybean yield has been significantly increased since the 1960s and is forecasted to reach 3.0 metric tonnes per hectare in 2028. It was observed that soybean yields in East Asian countries have been stagnant for some time. To meet the projected demands, it is important to consider breeding soybean cultivars with higher yield potentials and better tolerance to biotic and abiotic stresses. Wild soybean has been shown to provide an important resource for soybean breeding in terms of nutrition, biotic and abiotic stress tolerance, and yield-related traits (Table 3).
    Table 3. List of genes and QTLs found to be associated with specific traits using either wild soybean or a cross between wild and cultivated soybeans.
    Trait Gene/QTL Chromosome/Locus Group Source (accession) Reference
    Yield Related Traits
    Flower color Dihydroflavonol-4-reductase 4 (w4-s1 allele) (DFR) B2 (Ch 14) WS (CW13133) [46]
    Flavonoid 3′5′-hydroxylase (F3′5′H) and DFR B2 (Ch 14) WS (CW12700 and CW13381) [45]
    DFR2 (w4-s1) B2 (Ch 14) WS (CW13133) × CS (IT182932) [47]
    Oil and local breeding related traits QTLs 3, 8, 13, 17, WS × CS [17]
    Shoot fresh weight QTLs 6, 15, 19 WS (PI 483463) × CS (Hutcheson) [48]
    Biological nitrogen fixation related traits QTLs 7,8,12,17,18 WS (W05) × CS (C08) [49]
    Small seed, Yellow seed color,
    Soy sprout
        WS (PI135624) [50]
    High yield   B2 (U26) WS (PI 407305) [51]
    Yield, height and maturity QTLs C2, E, K and M WS × CS [52]
    Seed yield, 100-seed weight, seed filling period, maturity, height, lodging QTLs A1, J, N, H, F, L WS × CS [53]
      QTLs Multiple CS (NN86-4) × WS (PI342618B) [54]
      QTLs 1, 2, 6, 8, 13, 14, 17, 20 CS (Williams 82) × WS (PI 366121) [55]
    Biotic Stress Tolerance
    Soybean cyst nematode resistance Rhg1Rhg4 locus and QTLs G, A2, B1 CS (Magellan) × WS (PI 404198A) [56]
    Glyma18g196200, Glyma18g077900, Glyma18g078000, Glyma18g106800, Glyma18g107000, Glyma18g107100 Ch18 WS [57]
    Glyma18g106800, Glyma18g064100
    QTLs (cqSCN-006, cqSCN-007) I, J, K, O WS (PI 468916, 464925B) [58][59]
    Markers (A245_1 and Satt598) 15, 18 WS (PI 468916) [60]
    Resistance to southern root knot nematode     WS [61]
    Foxglove aphid resistance
    (Aulacorthum solani)
    Raso2 7 CS (Williams 82) × CS (PI 366121) [62]
    Aphid resistance (Aphis glycines) Rag3c and Rag6 8, 16 WS (85-32) [63]
    QTLs 8, 16, WS
    WS (PI 518282)
    [64]
    [65]
    Sclerotinia stem rot QTLs 3, 8, 20 WS × CS [66][67][68]
    Resistance to Mung bean yellow mosaic India virus   Ch 8 and 14 WS (PI 393551) [69]
    Resistance to Phtophthora soja QTLs J, I, G (Ch 16, 18, 20) WS (PI 407162) × CS (V71-370) [70]
    Abiotic Stress Tolerance
    Salt tolerance     WS (N23232 BB52) [71]
    Allele (Ncl locus)   WS (PI 483463) [72]
    QTLs D2 (Ch 17) WS (JWS156-1) [73]
    QTLs 3 WS [74]
    RLKs CaMs, JA/SA Signaling genes, MAPKs, WRKYs     [67][68]
    Tolerance to herbicide metribuzin     WS (PI 245331, PI 163453) [75]
    Root traits QTLs 8, 12 WS (PI 326582A) × CS (PI 552538) [76]
    Root architecture Glyma15g42220, Glyma06g46210, Glyma06g45910, Glyma06g45920, Glyma07g32480 6, 7, 15 WS (PI 407162) × CS (V71-370) [77]
    Alkalinity tolerance ALMT, LEA, ABC transporter, GLR, NRT/POT and SLAH genes Multiple WS (N24852) [78]
    Drought tolerance GsWRKY20     [79]
    Nutrition
    Seed protein content Marker pA-245 C CS (A81-356022) × WS (PI 468916) [80]
    Seed saturated fatty acid contents SNPs   GS [81]
    Glyma14g121400, Glyma16g068500 14, 16
    Seed unsaturated fatty acids Glyma16g014000, Glyma07g112100 7, 16
    Linolenic acid QTLs Multiple WS × CS [82]
    Pearl Japanese fermented product (Natto)     WS [61]
    Protein, oil, palmitic acid, stearic acid, oleic acid, linoleic acid, lenolenic acid QTLs 2, 7, 14, 16 WS [81]
    Saponin A Sg-1 Ch 7 WS [65][83]
    Sg-5 (Glyma15g39090) Ch 15 WS × CS [84][69]
    Glyma15g39090 Ch 15 WS [36]
    Seed antioxidant, phenolics, and flavonoids GmMATE1,2,4 Ch 18,19 WS (W05) × CS (C08) [85]
    So far, only a few characteristics of wild soybean contained in the CWLGC have been explored. Phenotypic screening towards biotic and abiotic resistance under diverse climate conditions would reveal more relevant results to meet current needs. In this regard, the CWLGC G. soja collection has been initially explored for the diversity in root system architecture (Figure 3b). Regarding biotic stress tolerance, no screening has yet been done, leaving the opportunity open for researchers interested in disease and insect pest resistance. Soybean cyst nematode, soybean mosaic virus, bacterial blight and wilt, and a whole list of fungal diseases could be the initial screening targets. Furthermore, early emergence and vigor traits in wild soybean have not yet been examined. Wild soybean seeds can survive in extreme climatic conditions in their native habitats, making them a good target for studying dormancy as well as longevity traits. These traits are important in terms of soybean improvement as well as ex-situ germplasm conservation. By analyzing the morphological variations in leaf shape in the CWLGC G. soja accessions, we can potentially unveil the process of domestication of this trait and enhance our understanding of the regulation of photosynthesis. A comparative and in-depth genome resequencing of G. soja chloroplast genomes could enhance our understanding of the domestication process of G. max, as well as of the traits regulated by the plastid genome. Up until now, only limited knowledge of these aspects is available [86][87].

    The entry is from 10.3390/agronomy10020214

    References

    1. Kofsky, J.; Zhang, H.; Song, B.-H. The untapped genetic reservoir: the past, current, and future applications of the wild soybean (Glycine soja). Front. Plant Sci. 2018, 9.
    2. Nawaz, M.A.; Rehman, H.M.; Imtiaz, M.; Baloch, F.S.; Lee, J.D.; Yang, S.H.; Lee, S.I.; Chung, G. Systems Identification and Characterization of Cell Wall Reassembly and Degradation Related Genes in Glycine max (L.) Merill, a Bioenergy Legume. Sci. Rep. 2017, 7, 10862.
    3. Lam, H.-M.; Remais, J.; Fung, M.-C.; Xu, L.; Sun, S.S.-M. Food supply and food safety issues in China. The Lancet 2013, 381, 2044–2053.
    4. Considine, M.J.; Siddique, K.H.; Foyer, C.H. Nature’s pulse power: Legumes, food security and climate change. J. Exp. Bot. 2017, 68, 1815–1818.
    5. Allender, C. The Second Report on the State of the World’s Plant. Genetic Resources for Food and Agriculture; FAO Commission on Genetic Resources for Food and Agriculture: Rome, Italy, 2010.
    6. Lin, B.B. Resilience in agriculture through crop diversification: adaptive management for environmental change. BioScience 2011, 61, 183–193.
    7. Hyten, D.L.; Song, Q.; Zhu, Y.; Choi, I.-Y.; Nelson, R.L.; Costa, J.M.; Specht, J.E.; Shoemaker, R.C.; Cregan, P.B. Impacts of genetic bottlenecks on soybean genome diversity. Proc. Natl. Acad. Sci. USA 2006, 103, 16666–16671.
    8. Cowling, W.; Li, L.; Siddique, K.; Henryon, M.; Berg, P.; Banks, R.; Kinghorn, B. Evolving gene banks: Improving diverse populations of crop and exotic germplasm with optimal contribution selection. J. Exp. Bot. 2016, 68, 1927–1939.
    9. Foyer, C.H.; Lam, H.-M.; Nguyen, H.T.; Siddique, K.H.; Varshney, R.K.; Colmer, T.D.; Cowling, W.; Bramley, H.; Mori, T.A.; Hodgson, J.M. Neglecting legumes has compromised human health and sustainable food production. Nat. Plant. 2016, 2, 16112.
    10. Muñoz, N.; Liu, A.; Kan, L.; Li, M.-W.; Lam, H.-M. Potential uses of wild germplasms of grain legumes for crop improvement. Int. J. Mol. Sci. 2017, 18, 328.
    11. Chung, G.; Singh, R.J. Broadening the genetic base of soybean: A multidisciplinary approach. Crit. Rev. Plant Sci. 2008, 27, 295–341.
    12. Ladizinsky, G.; Newell, C.; Hymowitz, T. Wide crosses in soybeans: Prospects and limitations. Euphytica 1979, 28, 421–423.
    13. Singh, R.; Nelson, R.L. Methodology for creating alloplasmic soybean lines by using Glycine tomentella as a maternal parent. Plant Breed. 2014, 133, 624–631.
    14. Singh, R.J. Cytogenetics and genetic introgression from wild relatives in soybean. Nucleus 2019, 62, 3–14.
    15. Singh, R.; Hymowitz, T. The genomic relationship between Glycine max (L.) Merr. and G. soja Sieb. and Zucc. as revealed by pachytene chromosome analysis. Theor. Appl. Genet. 1988, 76, 705–711.
    16. He, S.-L.; Wang, Y.-S.; Li, D.-Z.; Yi, T.-S. Environmental and historical determinants of patterns of genetic differentiation in wild soybean (Glycine soja Sieb. et Zucc). Sci. Rep. 2016, 6, 22795.
    17. Zhou, Z.; Jiang, Y.; Wang, Z.; Gou, Z.; Lyu, J.; Li, W.; Yu, Y.; Shu, L.; Zhao, Y.; Ma, Y. Resequencing 302 wild and cultivated accessions identifies genes related to domestication and improvement in soybean. Nat. Biotechnol. 2015, 33, 408–414.
    18. Wang, L.-x.; Lin, F.-y.; LI, L.-h.; Wei, L.; Zhe, Y.; LUAN, W.-j.; PIAO, R.-h.; Yuan, G.; NING, X.-c.; Li, Z. Genetic diversity center of cultivated soybean (Glycine max) in China–New insight and evidence for the diversity center of Chinese cultivated soybean. J. Integr. Agric. 2016, 15, 2481–2487.
    19. Leamy, L.J.; Lee, C.R.; Song, Q.; Mujacic, I.; Luo, Y.; Chen, C.Y.; Li, C.; Kjemtrup, S.; Song, B.H. Environmental versus geographical effects on genomic variation in wild soybean (Glycine soja) across its native range in northeast Asia. Ecol. Evolut. 2016, 6, 6332–6344.
    20. Hay, F.R.; Probert, R.J. Advances in seed conservation of wild plant species: A review of recent research. Conserv. Physiol. 2013, 1.
    21. Castañeda-Álvarez, N.P.; Khoury, C.K.; Achicanoy, H.A.; Bernau, V.; Dempewolf, H.; Eastwood, R.J.; Guarino, L.; Harker, R.H.; Jarvis, A.; Struik, P.C.; et al. Global conservation priorities for crop wild relatives. Nat. Plant. 2016, 2, 16022.
    22. Lee, J.D.; Shannon, J.; Vuong, T.; Moon, H.; Nguyen, H.; Tsukamoto, C.; Chung, G. Genetic diversity in wild soybean (Glycine soja Sieb. and Zucc.) accessions from southern islands of Korean peninsula. Plant Breed. 2010, 129, 257–263.
    23. Nawaz, M.A.; Yang, S.H.; Rehman, H.M.; Baloch, F.S.; Lee, J.D.; Park, J.H.; Chung, G. Genetic diversity and population structure of Korean wild soybean (Glycine soja Sieb. and Zucc.) inferred from microsatellite markers. Biochem. Syst. Ecol. 2017, 71, 87–96.
    24. Carter, T.; Nelson, R.; Sneller, C.; Cui, Z. Genetic diversity in soybean. Soybean Monogr. Am. Soc. Agron. Madison Wis. USA 2004.
    25. Li, M.-W.; Wang, Z.; Jiang, B.; Kaga, A.; Wong, F.-L.; Zhang, G.; Han, T.; Chung, G.; Nguyen, H.; Lam, H.-M. Impacts of genomic research on soybean improvement in East Asia. Theor. Appl. Genet. 2019, 1–24.
    26. Lee, G.-A.; Crawford, G.W.; Liu, L.; Sasaki, Y.; Chen, X. Archaeological soybean (Glycine max) in East Asia: Does size matter? PLoS ONE 2011, 6, e26720.
    27. Crawford, G.W.; Lee, G.-A. Agricultural origins in the Korean Peninsula. Antiquity 2003, 77, 87.
    28. Barnes, G.L. Archaeology of East. Asia: The Rise of Civilization in China, Korea and Japan; Oxbow Books: Oxford, UK; Casemate Publishing: Pennsylvania, PA, USA, 2015.
    29. Maxted, N.; Kell, S. Establishment of a global network for the in situ conservation of crop wild relatives: Status and needs. FAO Comm. Genet. Resour. Food Agric. Rome 2009, 266, 509.
    30. Kim, M.Y.; Van, K.; Kang, Y.J.; Kim, K.H.; Lee, S.-H. Tracing soybean domestication history: From nucleotide to genome. Breed. Sci. 2012, 61, 445–452.
    31. Lee, S.; Kim, J.-H.; Sundaramoorthy, J.; Park, G.T.; Lee, J.-D.; Kim, J.H.; Chung, G.; Seo, H.S.; Song, J.T. Identification of GmSALT3 haplotypes and development of molecular markers based on their diversity associated with salt tolerance in soybean. Mol. Breed. 2018, 38, 86.
    32. Panneerselvam, K.; Tsukamoto, C.; Honda, N.; Kikuchi, A.; Lee, J.D.; Yang, S.H.; Chung, G. Saponin polymorphism in the Korean wild soybean (Glycine soja Sieb. and Zucc.). Plant Breed. 2013, 132, 121–126.
    33. Krishnamurthy, P.; Lee, J.M.; Tsukamoto, C.; Takahashi, Y.; Singh, R.J.; Lee, J.D.; Chung, G. Evaluation of genetic structure of Korean wild soybean (Glycine soja) based on saponin allele polymorphism. Genet. Res. Crop Evol. 2014, 61, 1121–1130.
    34. Krishnamurthy, P.; Tsukamoto, C.; Singh, R.J.; Lee, J.-D.; Kim, H.-S.; Yang, S.-H.; Chung, G. The Sg-6 saponins, new components in wild soybean (Glycine soja Sieb. and Zucc.): polymorphism, geographical distribution and inheritance. Euphytica 2014, 198, 413–424.
    35. Krishnamurthy, P.; Tsukamoto, C.; Takahashi, Y.; Hongo, Y.; Singh, R.J.; Lee, J.D.; Chung, G. Comparison of saponin composition and content in wild soybean (Glycine soja Sieb. and Zucc.) before and after germination. Biosci. Biotechnol. Biochem. 2014, 78, 1988–1996.
    36. Rehman, H.M.; Nawaz, M.A.; Shah, Z.H.; Yang, S.H.; Chung, G. Functional characterization of naturally occurring wild soybean mutant (sg-5) lacking astringent saponins using whole genome sequencing approach. Plant Sci. 2017.
    37. Sundaramoorthy, J.; Park, G.T.; Mukaiyama, K.; Tsukamoto, C.; Chang, J.H.; Lee, J.-D.; Kim, J.H.; Seo, H.S.; Song, J.T. Molecular elucidation of a new allelic variation at the Sg-5 gene associated with the absence of group A saponins in wild soybean. PLoS ONE 2018, 13, e0192150.
    38. Krishnamurthy, P.; Lee, J.D.; Ha, B.K.; Chae, J.H.; Song, J.T.; Tsukamoto, C.; Singh, R.J.; Chung, G. Genetic characterization of group A acetylsaponin-deficient mutants from wild soybean (Glycine soja Sieb. and Zucc.). Plant Breed. 2015, 134, 316–321.
    39. Rehman, H.M.; Nawaz, M.A.; Shah, Z.H.; Chung, G.; Yang, S.H. Molecular Elucidation of Two Novel Seed Specific Flavonoid Glycosyltransferases in Soybean. J. Plant Biol. 2018, 61, 320–329.
    40. Krishnamurthy, P.; Singh, R.J.; Tsukamoto, C.; Park, J.H.; Lee, J.D.; Chung, G. Kunitz trypsin inhibitor polymorphism in the Korean wild soybean (Glycine soja Sieb. & Zucc.). Plant Breed. 2013, 132, 311–316.
    41. Nawaz, M.A.; Golokhvast, K.S.; Rehman, H.M.; Tsukamoto, C.; Kim, H.-S.; Yang, S.H.; Chung, G. Soyisoflavone diversity in wild soybeans (Glycine soja Sieb. & Zucc.) from the main centres of diversity. Biochem. Syst. Ecol. 2018, 77, 16–21.
    42. Tsukamoto, C.; Nawaz, M.A.; Kurosaka, A.; Le, B.; Lee, J.D.; Son, E.; Yang, S.H.; Kurt, C.; BALOCH, F.S.; Chung, G. Isoflavone profile diversity in Korean wild soybeans (Glycine soja Sieb. & Zucc.). Turk. J. Agric. For. 2018, 42, 248–261.
    43. Chae, J.-H.; Ha, B.-K.; Chung, G.; Park, J.-E.; Park, E.; Ko, J.-M.; Shannon, J.G.; Song, J.T.; Lee, J.-D. Identification of environmentally stable wild soybean genotypes with high alpha-linolenic acid concentration. Crop Sci. 2015, 55, 1629–1636.
    44. Ali, M.; Krishnamurthy, P.; El-Hadary, M.; Kim, J.; Nawaz, M.; Yang, S.; Chung, G. Identification and expression profiling of a new β-amyrin synthase gene (GmBAS3) from soybean. Russ. J. Plant physiol. 2016, 63, 383–390.
    45. Sundaramoorthy, J.; Park, G.T.; Chang, J.H.; Lee, J.-D.; Kim, J.H.; Seo, H.S.; Chung, G.; Song, J.T. Identification and molecular analysis of four new alleles at the W1 locus associated with flower color in soybean. PLoS ONE 2016, 11, e0159865.
    46. Park, G.T.; Sundaramoorthy, J.; Lee, S.; Lee, J.-D.; Kim, J.H.; Park, S.-K.; Seo, H.S.; Chung, G.; Song, J.T. Color Variation in a Novel Glycine soja Mutant W4-S1 with Pinkish-White Flowers Is Controlled by a Single Recessive Allele at the W4 Locus. Crop Sci. 2017, 57, 3112–3121.
    47. Park, G.T.; Sundaramoorthy, J.; Lee, S.; Lee, J.-D.; Kim, J.H.; Park, S.-K.; Seo, H.S.; Chung, G.; Song, J.T. Color Variation in a Novel Mutant with Pinkish-White Flowers Is Controlled by a Single Recessive Allele at the Locus. Crop Sci. 2017, 57, 3112–3121.
    48. Asekova, S.; Kulkarni, K.P.; Patil, G.; Kim, M.; Song, J.T.; Nguyen, H.T.; Shannon, J.G.; Lee, J.-D. Genetic analysis of shoot fresh weight in a cross of wild (G. soja). Mol. Breed. 2016, 36, 1–15.
    49. Muñoz, N.; Qi, X.; Li, M.; Xie, M.; Gao, Y.; Cheung, M.; Wong, F.; Lam, H. Improvement in nitrogen fixation capacity could be part of the domestication process in soybean. Heredity 2016, 117, 84–93.
    50. Fehr, W.; Cianzio, S.; Welke, G. Registration of’SS202’soybean. Crop Sci. 1990, 30.
    51. Concibido, V.; La Vallee, B.; Mclaird, P.; Pineda, N.; Meyer, J.; Hummel, L.; Yang, J.; Wu, K.; Delannay, X. Introgression of a quantitative trait locus for yield from Glycine soja into commercial soybean cultivars. Theor. Appl. Genet. 2003, 106, 575–582.
    52. Wang, D.; Graef, G.; Procopiuk, A.; Diers, B. Identification of putative QTL that underlie yield in interspecific soybean backcross populations. Theor. Appl. Genet. 2004, 108, 458–467.
    53. Li, D.; Pfeiffer, T.; Cornelius, P. Soybean QTL for yield and yield components associated with alleles. Crop Sci. 2008, 48, 571–581.
    54. Wang, W.; Li, X.; Chen, S.; Song, S.; Gai, J.; Zhao, T. Using presence/absence variation markers to identify the QTL/allele system that confers the small seed trait in wild soybean (Glycine soja Sieb. & Zucc.). Euphytica 2016, 208, 101–111.
    55. Kulkarni, K.P.; Asekova, S.; Lee, D.-H.; Bilyeu, K.; Song, J.T.; Lee, J.-D. Mapping QTLs for 100-seed weight in an interspecific soybean cross of Williams 82 (Glycine max) and PI 366121 (Glycine soja). Crop Pasture Sci. 2017, 68, 148–155.
    56. Guo, B.; Sleper, D.A.; Nguyen, H.T.; Arelli, P.R.; Shannon, J.G. Quantitative trait loci underlying resistance to three soybean cyst nematode populations in soybean PI 404198A. Crop Sci. 2006, 46, 224–233.
    57. Zhang, H.; Li, C.; Davis, E.L.; Wang, J.; Griffin, J.D.; Kofsky, J.; Song, B.-H. Genome-wide association study of resistance to soybean cyst nematode (Heterodera glycines) HG Type 2.5. 7 in wild soybean (Glycine soja). Front. Plant Sci. 2016, 7, 1214.
    58. Winter, S.M.; Shelp, B.J.; Anderson, T.R.; Welacky, T.W.; Rajcan, I. QTL associated with horizontal resistance to soybean cyst nematode in Glycine soja PI464925B. Theor. Appl. Genet. 2007, 114, 461–472.
    59. Yu, N.; Diers, B.W. Fine mapping of the SCN resistance QTL cqSCN-006 and cqSCN-007 from Glycine soja PI 468916. Euphytica 2017, 213, 54.
    60. Wang, D.; Diers, B.W.; Arelli, P.; Shoemaker, R. Loci underlying resistance to race 3 of soybean cyst nematode in Glycine soja plant introduction 468916. Theor. Appl. Genet. 2001, 103, 561–566.
    61. Carter, T.E.; Huei, E.; Burton, J.; Farmer, F.; Gizlice, Z. Registration of ‘Pearl’soybean. Crop Sci. 1995, 35, 1713.
    62. Lee, J.S.; Yoo, M.-h.; Jung, J.K.; Bilyeu, K.D.; Lee, J.-D.; Kang, S. Detection of novel QTLs for foxglove aphid resistance in soybean. Theor. Appl. Genet. 2015, 128, 1481–1488.
    63. Zhang, S.; Zhang, Z.; Wen, Z.; Gu, C.; An, Y.-Q.C.; Bales, C.; DiFonzo, C.; Song, Q.; Wang, D. Fine mapping of the soybean aphid-resistance genes Rag6 and Rag3c from Glycine soja 85-32. Theor. Appl. Genet. 2017, 130, 2601–2615.
    64. Zhang, S.; Zhang, Z.; Bales, C.; Gu, C.; DiFonzo, C.; Li, M.; Song, Q.; Cregan, P.; Yang, Z.; Wang, D. Mapping novel aphid resistance QTL from wild soybean, Glycine soja 85-32. Theor. Appl. Genet. 2017, 130, 1941–1952.
    65. Hill, C.B.; Li, Y.; Hartman, G.L. Resistance of Glycine species and various cultivated legumes to the soybean aphid (Homoptera: Aphididae). J. Econ. Entomol. 2004, 97, 1071–1077.
    66. Iquira, E.; Humira, S.; François, B. Association mapping of QTLs for sclerotinia stem rot resistance in a collection of soybean plant introductions using a genotyping by sequencing (GBS) approach. BMC Plant Biol. 2015, 15, 5.
    67. Zhang, H.; Song, B.-H. RNA-seq data comparisons of wild soybean genotypes in response to soybean cyst nematode (Heterodera glycines). Genomics Data 2017, 14, 36–39.
    68. Zhang, H.; Song, Q.; Griffin, J.D.; Song, B.-H. Genetic architecture of wild soybean (Glycine soja) response to soybean cyst nematode (Heterodera glycines). Mol. Genet. Genomics 2017, 292, 1257–1265.
    69. Rani, A.; Kumar, V.; Gill, B.; Shukla, S.; Rathi, P.; Singh, R. Mapping of duplicate dominant genes for Mungbean yellow mosaic India virus resistance in Glycine soja. Crop Sci. 2018, 58, 1566–1574.
    70. Tucker, D.; Maroof, S.; Mideros, S.; Skoneczka, J.; Nabati, D.; Buss, G.; Hoeschele, I.; Tyler, B.; St Martin, S.; Dorrance, A. Mapping quantitative trait loci for partial resistance to Phytophthora sojae in a soybean interspecific cross. Crop Sci. 2010, 50, 628–635.
    71. Luo, Q.; Yu, B.; Liu, Y. Differential sensitivity to chloride and sodium ions in seedlings of Glycine max and G. soja under NaCl stress. J. Plant Physiol. 2005, 162, 1003–1012.
    72. Lee, J.-D.; Shannon, J.G.; Vuong, T.D.; Nguyen, H.T. Inheritance of salt tolerance in wild soybean (Glycine soja Sieb. and Zucc.) accession PI483463. J. Hered. 2009, 100, 798–801.
    73. Tuyen, D.; Lal, S.; Xu, D. Identification of a major QTL allele from wild soybean (Glycine soja Sieb. & Zucc.) for increasing alkaline salt tolerance in soybean. Theor. Appl. Genet. 2010, 121, 229–236.
    74. Ha, B.-K.; Vuong, T.D.; Velusamy, V.; Nguyen, H.T.; Shannon, J.G.; Lee, J.-D. Genetic mapping of quantitative trait loci conditioning salt tolerance in wild soybean (Glycine soja) PI 483463. Euphytica 2013, 193, 79–88.
    75. Kilen, T.C.; He, G. Identification and inheritance of metribuzin tolerance in wild soybean. Crop Sci. 1992, 32, 684–685.
    76. Manavalan, L.P.; Prince, S.J.; Musket, T.A.; Chaky, J.; Deshmukh, R.; Vuong, T.D.; Song, L.; Cregan, P.B.; Nelson, J.C.; Shannon, J.G. Identification of novel QTL governing root architectural traits in an interspecific soybean population. PLoS ONE 2015, 10, e0120490.
    77. Prince, S.J.; Song, L.; Qiu, D.; dos Santos, J.V.M.; Chai, C.; Joshi, T.; Patil, G.; Valliyodan, B.; Vuong, T.D.; Murphy, M. Genetic variants in root architecture-related genes in a Glycine soja accession, a potential resource to improve cultivated soybean. BMC Genomics 2015, 16, 132.
    78. Zhang, J.; Wang, J.; Jiang, W.; Liu, J.; Yang, S.; Gai, J.; Li, Y. Identification and analysis of NaHCO3 stress responsive genes in wild soybean (Glycine soja) roots by RNA-seq. Front. Plant Sci. 2016, 7, 1842.
    79. Ning, W.; Zhai, H.; Yu, J.; Liang, S.; Yang, X.; Xing, X.; Huo, J.; Pang, T.; Yang, Y.; Bai, X. Overexpression of Glycine soja WRKY20 enhances drought tolerance and improves plant yields under drought stress in transgenic soybean. Mol. Breed. 2017, 37, 19.
    80. Diers, B.W.; Keim, P.; Fehr, W.; Shoemaker, R. RFLP analysis of soybean seed protein and oil content. Theor. Appl. Genet. 1992, 83, 608–612.
    81. Leamy, L.J.; Zhang, H.; Li, C.; Chen, C.Y.; Song, B.-H. A genome-wide association study of seed composition traits in wild soybean (Glycine soja). BMC Genomics 2017, 18, 18.
    82. Pantalone, V.; Rebetzke, G.; Burton, J.; Wilson, R. Genetic regulation of linolenic acid concentration in wild soybean Glycine soja accessions. J. Am. Oil Chem. Soc. 1997, 74, 159–163.
    83. Park, J.; Kim, J.H.; Krishnamurthy, P.; Tsukamoto, C.; Song, J.T.; Chung, G.; Shannon, J.G.; Lee, J.-D. Characterization of a New Allele of the Saponin-Synthesizing Gene in Soybean. Crop Sci. 2016, 56, 385–391.
    84. Yano, R.; Takagi, K.; Takada, Y.; Mukaiyama, K.; Tsukamoto, C.; Sayama, T.; Kaga, A.; Anai, T.; Sawai, S.; Ohyama, K. Metabolic switching of astringent and beneficial triterpenoid saponins in soybean is achieved by a loss-of-function mutation in cytochrome P450 72A69. Plant J. 2017, 89, 527–539.
    85. Li, M.-W.; Muñoz, N.B.; Wong, C.-F.; Wong, F.-L.; Wong, K.-S.; Wong, J.W.-H.; Qi, X.; Li, K.-P.; Ng, M.-S.; Lam, H.-M. QTLs regulating the contents of antioxidants, phenolics, and flavonoids in soybean seeds share a common genomic region. Front. Plant Sci. 2016, 7.
    86. Fang, C.; Ma, Y.; Yuan, L.; Wang, Z.; Yang, R.; Zhou, Z.; Liu, T.; Tian, Z. Chloroplast DNA Underwent Independent Selection from Nuclear Genes during Soybean Domestication and Improvement. J. Genet. Genomics = Yi Chuan Xue Bao 2016, 43, 217.
    87. Asaf, S.; Khan, A.L.; Khan, M.A.; Imran, Q.M.; Kang, S.-M.; Al-Hosni, K.; Jeong, E.J.; Lee, K.E.; Lee, I.-J. Comparative analysis of complete plastid genomes from wild soybean (Glycine soja) and nine other Glycine species. PLoS ONE 2017, 12, e0182281.
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