Transgressive inheritance of broad-spectrum resistance to Heterodera glycines and Meloidogyne incognita in soybean chromosome segment substitution lines

Transgressive inheritance of broad-spectrum resistance to Heterodera glycines and Meloidogyne incognita in soybean chromosome segment substitution lines | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Transgressive inheritance of broad-spectrum resistance to Heterodera glycines and Meloidogyne incognita in soybean chromosome segment substitution lines Minghui Huang, Chunyan Liu, Lei Zhao, Yanan Zhao, Ye Jiang, Doudou Chang, and 3 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7587031/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Soybean cyst nematode ( Heterodera glycines , SCN) and southern root-knot nematode ( Meloidogyne incognita , RKN) are major limitation to soybean production, with few resistance sources available. Using 199 and 177 chromosome segment substitution lines (CSSLs) derived from Glycine max cv. Suinong14 and G. soja accession ZYD00006, we evaluated resistance to SCN HG type 1.2.3.5.6.7 (race 4) and RKN race 1, and compared the loci with those previously mapped for SCN HG type 2.5.7 (race 5). Despite both parents being susceptible, many CSSLs exhibited resistance levels exceeding both parents, while others showed extreme susceptibility, demonstrating strong transgressive segregation. QTL mapping identified 6 FI-associated and 9 CGR-associated loci for SCN race 4, and 17–25 significant loci for RKN across two inoculum levels. Single resistance loci generally conferred only moderate effects, but the combination of multiple favorable alleles from both parents produced higher resistance. Comparative analysis revealed several shared QTL intervals among SCN races 4 and 5 and RKN race 1, suggesting partial common genetic control. These findings demonstrate that pyramiding multiple loci from wild and cultivated soybean can generate transgressive inheritance, producing resistant lines beyond parental performance and offering valuable targets for breeding durable, broad-spectrum nematode resistance. Glycine max G. soja Soybean cyst nematode Root-knot nematode Transgressive inheritance Chromosome segment substitution lines QTL mapping Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Introduction Soybean ( Glycine max L. Merr.) is a globally important oilseed crop and a key component of food and agricultural systems. China ranks as the fourth-largest soybean producer, following the United States, Brazil, and Argentina. Among the major biotic constraints, plant-parasitic nematodes (PPNs) are the second most damaging group of pathogens globally, significantly threatening food security and crop yields. Among PPNs, the soybean cyst nematode ( Heterodera glycines , SCN) and root-knot nematodes ( Meloidogyne spp., RKN) are the most prevalent and destructive nematodes, causing serious yield losses across diverse cropping systems (Bent et al. 2022; Thakur et al. 2024 ). Both SCN and RKN are obligate sedentary endoparasites. Their infective second-stage juveniles (J2s) invade roots and secrete effectors through their stylets to establish specialized feeding sites — syncytia in SCN and giant cells in RKN — disrupting nutrient flow and reducing yield (Niblack et al. 2006 ; Mitchum and Baum 2008 ). In China, SCN is especially problematic in the Northeast, the country’s primary soybean-producing region. Yield losses typically range from 20–30%, rising to 60–70% under severe infestations (Peng 2021). Annual losses due to SCN exceed $ 1 billion in the U.S. (Koenning and Wrather 2006 ) and 600 million yuan in China (Peng 2021). RKNs, particularly Meloidogyne incognita , infect over 5,500 plant species and cause global crop losses of approximately $ 7.8 billion (Caillaud et al. 2008 ; Nicol et al. 2011 ). In China, RKN-induced yield reductions in vegetables can exceed 30%, with annual economic losses surpassing 3 billion yuan (Peng 2021). M. incognita is the dominant RKN species worldwide and in China. Although historically confined to central regions, it is now spreading to northern provinces via protected cultivation systems. Greenhouse conditions (~ 23°C) promote overwintering and rapid reproduction (Zhao et al. 2011 ; Hong et al. 2018 ), accelerating its spread in regions such as Liaoning and Heilongjiang and increasing its threat to soybean production. Current control strategies for SCN and RKN rely primarily on resistant cultivars and crop rotation. However, both H. glycines and M. incognita exhibit high genetic diversity and complex race structures (HG types and pathotypes), making breeding efforts challenging. Most commercial SCN-resistant cultivars are derived from only two sources (Peking and PI 88788), leading to the emergence of virulent nematode populations, particularly in Heilongjiang, where dominant SCN races have shifted from race 3 to more aggressive races 4 and 14 (Acharya et al. 2016 ; Bayless et al. 2018 ; Huang et al. 2022a ; You et al. 2024 ). Although M. incognita has been considered a warm-climate pest, its rapid expansion in protected environments highlights the urgency of addressing its threat in colder regions, including Northeast China (Zhao et al. 2010 ; Li et al. 2016 ). While international studies dominate RKN resistance research (Schmitt and Bellé 2016; Canella et al. 2022; Gontijio et al. 2024), domestic progress is increasing. For example, Li et al. ( 2016 ) screened native soybean germplasm under greenhouse conditions in Heilongjiang for M. incognita resistance. SCN resistance is typically assessed using the Female Index (FI), while the cysts per gram of root (CGR) provides a root-weight-adjusted metric (Huang et al. 2021 ). In contrast, no standardized system exists for M. incognita resistance evaluation. Most studies use the Gall Index (GI), often without accounting for egg counts or standardized inoculum, which hampers precision. Thus, developing a robust and comprehensive evaluation framework for RKN resistance remains a key priority. Resistance to both SCN and RKN in soybean is polygenic and quantitatively inherited (Concibido et al. 2004 ;Li et al. 2018 ; Vieira et al. 2022 ; Bent et al. 2022). To date, over 300 quantitative trait loci (QTLs) associated with SCN resistance have been mapped across all 20 soybean chromosomes (SoyBase, www.soybase.org ), reflecting the complex genetic architecture of resistance. Among them, two major loci — rhg1 on chromosome 18 and Rhg4 on chromosome 8, have been most extensively characterized (Cook et al. 2012 , 2014 ; Liu et al. 2012 , 2017 ). The rhg1 locus comprises multiple genes, including α-SNAP, amino acid transporter (AAT), and WI12, as well as other SNAP family members such as GmSNAP11 and GmSNAP02 , which play pivotal roles in nematode resistance (Bayless et al. 2016 , 2019 ; Guo et al. 2019 ; Dong et al. 2022; Han et al. 2023 ; Usovsky et al. 2023 ; Du et al. 2024). In contrast, Rhg4 encodes a serine hydroxymethyltransferase ( GmSHMT08 ), and its resistance effect is influenced by copy number variation and allelic dosage at both rhg1 and Rhg4 loci (Patil et al. 2019 ). Despite this progress, additional minor and environment-sensitive QTLs continue to be discovered, underscoring the need for deeper exploration of resistance loci, particularly in wild-derived populations and under field-relevant pathogen pressures. However, resistance QTLs for M. incognita are less well defined. Early mapping in PI 96354 identified two QTLs on Chr10 and Chr18 (Tamulonis et al. 1997 ), with later studies reporting additional loci on Chr6, 7, 8, 13, and 17 (Ha et al. 2004 ; Fourie et al. 2008 ; Xu et al. 2013 ; Jiao et al. 2015 ; Vuong et al. 2021 ). Li et al. ( 2018 ) reported that four major QTLs (Chr 10, 18, 20, and 7) associated with M. incognita resistance. A recent genome-wide association study (GWAS) involving 717 accessions detected major associations on Chr10 and minor loci on Chr11 (Vieira et al. 2022 ). However, many of these intervals are large and based solely on GI, limiting their practical utility. To improve QTL resolution, high-density genetic maps and multidimensional resistance indices are essential. Despite identification of major resistance loci, novel and durable sources of resistance are urgently required to enhance breeding pipelines. Traditional QTL mapping populations (e.g., F ₂ , RILs, DH) often suffer from broad confidence intervals due to genetic background noise. Chromosome Segment Substitution Lines (CSSLs), developed by repeated backcrossing and selfing, enable precise QTL mapping in near-isogenic backgrounds. By substituting specific genomic segments from a donor parent into a recurrent background, CSSLs improve the detection of both major and minor QTLs (Balakrishnan et al. 2019 ). The CSSL population used in this study, derived from Suinong14 (recurrent parent) and ZYD00006 (wild soybean donor), has been under development in Northeast China for over a decade (Xin et al. 2016 ; Jiang et al. 2020 ; Wang et al. 2023 ; Jong et al. 2024 ). This population has previously been applied to map QTLs for seed quality, nodulation, drought, cold tolerance, and more recently, transgressive resistance to H. glycines HG Type 2.5.7 (race 5), based on FI and CGR (Huang et al. 2021 ). However, that study did not address virulent H. glycines race 4 or the root-knot nematode M. incognita , which are emerging threats to soybean production and require further investigation. In this study, we aimed to (1) assess the resistance of the Suinong14 × ZYD00006 CSSL population to H. glycines race 4 and M. incognita race 1 under different inoculation pressures, (2) identify novel QTLs associated with resistance traits, and (3) evaluate transgressive segregation patterns and potential elite lines for breeding. Resistance was evaluated using multiple indices: FI and CGR for soybean cyst nematode, and GI, reproduction factor (RF), egg masses per gram root (EMR), and eggs per gram root (EGR) for root-knot nematode. High-resolution QTL mapping was conducted using a published physical map (Huang et al. 2021 ). These efforts will deepen our understanding of the genetic architecture of nematode resistance and support the development of soybean cultivars with durable, broad-spectrum resistance suited to the agroecological conditions of Northeast China. Materials and methods Plant and nematode materials Soybean chromosome segment substitution lines (CSSLs) at the BC 3 F 7 -BC 8 F 2 generations were developed through backcrossing between the recurrent parent G. max ‘Suinong14’ and the wild donor parent G. soja ‘ZYD00006’ (Huang et al. 2021; Li et al. 2021; Pan et al. 2022; Wang et al. 2023; Jong et al. 2024). The CSSL population, along with both parental lines, was obtained from Northeast Agricultural University. For resistance screening, 199 CSSLs were evaluated against H. glycines HG type 1.2.3.5.6.7 (designated as SCN race 4), and 177 CSSLs were evaluated against M. incognita (RKN race 1). Susceptible soybean cultivars ‘Dongsheng 1’ and ‘Lee68’ were used as propagation hosts and resistance controls for SCN, respectively, while M. incognita was maintained on tomato ( Solanum lycopersicum ) cultivar ‘Zhongshu 4’. Nematode cultures were maintained in a greenhouse under controlled conditions (22-28 °C, 16 h light/8 h dark photoperiod) at the Northeast Institute of Geography and Agroecology, Chinese Academy of Sciences. Preparation of SCN and RKN inocula SCN cysts were extracted from infected soybean roots and surrounding soil 35-40 days post-inoculation. Cysts were crushed to release eggs, and second-stage juveniles (J2s) were hatched in sterile water containing 3 mM ZnSO₄ at 28 °C. For RKN, roots were harvested 42 days post-inoculation, washed thoroughly, and immersed in 50 mL of 1% sodium hypochlorite (NaClO) solution, then shaken either on a shaker or by hand for 3 minutes. The suspension was filtered through a nested sieve set (75 μm upper, 25 μm lower) and rinsed with pressurized water. Eggs retained on the lower sieve were transferred for hatching as described for SCN. J2s and eggs were counted under a stereo microscope (Olympus, Japan) for subsequent inoculation. Resistance evaluation of soybean to SCN and RKN Seeds were sown in 8 cm × 12 cm pots filled with sterilized soil and sand (1:1 v/v). After emergence, seedlings were thinned to one per pot. At 8-9 days after planting, two 4-5 cm-deep holes were made near each seedling for nematode inoculation. Each plant was inoculated with either 1,000 SCN J2s or 5,000/10,000 RKN eggs. SCN resistance was assessed 21 days post-inoculation by counting cysts on washed roots under fluorescent light. For RKN, roots were harvested 42 days post-inoculation, dried, and stained in 75 mg/L erioglaucine for 30 min. Egg masses were counted under a light microscope, and fresh root weights were recorded. Eggs were extracted as described in Section 2.2. All experiments used a randomized complete block design with 5-6 biological replicates per line and were repeated twice. RKN evaluations were conducted at two inoculation doses, and ‘Lee68’ served as a susceptible control in all experiments. SCN and RKN resistance evaluation SCN resistance was quantified using the FI, calculated as the mean cyst count on a test line divided by the mean count on the susceptible control ‘Lee68’, multiplied by 100. Classification followed Schmitt and Shannon (1992): FI 1-10 = resistant, 11-30 = moderately resistant, 31-60 = moderately susceptible, > 60 = susceptible. An additional index, CGR was calculated by dividing cyst number by root weight to adjust for biomass. For RKN, four indices were used: GI, RF, EMR and EGR. EMR was calculated by dividing egg mass number by root weight, and EGR by dividing egg number by root weight. RF was calculated as total eggs per plant divided by eggs inoculated. Resistance levels based on EGR were: highly resistant (< 1,000), resistant (1,000 - 4,999), susceptible (5,000 - 9,999), and highly susceptible (≥ 10,000). QTL detection for SCN and RKN resistance Quantitative trait loci (QTLs) for SCN (FI and CGR) and RKN (GI, RF, EMR, EGR) resistance were identified using MapQTL 5.0 (Van Ooijen 2004) with phenotypic data and a SNP-based physical map comprising 3,780 bin markers (Huang et al. 2021). The Kruskal-Wallis (K*) test and multiple QTL mapping (MQM) were applied. Significant QTLs were defined by P 2 in MQM. LOD thresholds were determined by 1,000 permutations at α = 0.05 (Churchill and Doerge 1994). QTLs with P < 0.1 were visualized using MapChart 2.3 (Voorrips 2002). QTLs were named following Huang et al. (2021). SCN QTLs were denoted as Hg4-Gs/GmXXa/b-FI/CGR , and RKN QTLs as Mi1-Gs/GmXXa/b-GI/RF/EM/EGR . Here, ‘ Hg 4’ and ‘ Mi 1’ refer to H. glycines race 4 and M. incognita race 1, ‘ Gs ’ or ‘ Gm ’ indicate the allele origin ( G. soja or G. max ), ‘XX’ the chromosome number, and ‘a/b’ distinguish multiple QTLs on the same chromosome. Statistical analyses Phenotypic data were analyzed by one-way ANOVA using IBM SPSS Statistics 26.0. Tukey’s HSD test ( P < 0.05) was used to compare cyst and FI values among lines. Correlations between resistance parameters were evaluated using Microsoft Excel 2013. Results Phenotypic identification of CSSLs and p arents to H. glycines HG type 1.2.3.5.6.7 Phenotypic evaluation of CSSLs and their parental lines against H. glycines HG type 1.2.3.5.6.7 (SCN race 4) revealed no significant difference in cysts/plant and FI among the parent Suinong14 (127.4 ± 0.94; 105.11 ± 9.03), ZYD00006 (108 ± 9.75; 89.11 ± 8.05), and the susceptible control Lee68 (122.1 ± 13.92; 100 ± 11.48) (Fig. 1, Table S1). However, G. soja ZYD00006 exhibited a markedly higher number of CGR (113.52 ± 17.29) compared to G. max Suinong14 (50.11 ± 4.75) and the susceptible cultivar Lee68 (43.29 ± 7.95). Among the 199 CSSLs evaluated, FI values ranged from 45.87 to 164.05, and CGR values ranged from 24.63 to 118.07, indicating considerable phenotypic diversity. Several CSSLs displayed reduced susceptibility compared to both parents. Notably, Lines 205 and 193 showed the lowest FI (45.87 ± 12.08; 58.17 ± 33.42) and CGR (36.07 ± 9.23; 24.63 ± 13.91), with reductions of 48.5% and 50.9%, respectively, compared with ZYD00006 and Suinong14. Forty-four CSSLs had CGR lower than Suinong14, representing 22.1% transgressive resistance. While no line had FI < 30 (highly resistant threshold), nearly half (49.8%) of CSSLs exhibited FI values lower than Suinong14 (FI = 105.11 ± 9.03), indicating moderate resistance improvements. Eight CSSLs (Lines 4, 24, 50, 57, 85, 92, 193, and 205) combined CGR < 60 with FI < 70, identifying them as promising candidates for SCN resistance breeding. Some lines displayed incongruent FI and CGR values (e.g., Line 213: CGR = 44.94 ± 3.84; FI = 103.41 ± 8.43), suggesting different loci may control these traits. These results indicate that G. soja introgressions can enhance SCN resistance, even in the absence of highly resistant parents, and that selected CSSLs may serve as valuable pre-breeding resources under high disease pressure. Evaluation of CSSLs under different inoculum levels of M. incongnita Phenotypic responses of CSSLs to M. incognita race 1 at 5,000 and 10,000 eggs per plant are summarized in Fig. 2 and Table S1. No significant differences were observed between Suinong14 and ZYD00006 for GI, RF, EMR, or EGR at either inoculum level. At 5,000 eggs per plant, most CSSLs performed better than Suinong14: 40 lines had reduced GI, 6 had lower RF, 135 had decreased EMR, and 161 had reduced EGR. Many lines were highly resistant, with minimal nematode reproduction. At 10,000 eggs per plant, transgressive resistance remained: 45% of lines had increased GI, 33% reduced RF, 37% reduced EMR, and 43% reduced EGR. However, the proportion of resistant lines (EGR < 5,000) decreased by 21.47% relative to the lower inoculum, indicating dose-dependent declines in resistance. Several CSSLs, including Lines 206, 210, 213, and 214, maintained stable resistance across doses, and eleven lines (e.g., 46, 119, 152, 154, 155, 156, 158, 159, 175, 189, 190) had EGR between 1,000-2,000 at high dose and < 1,000 at low dose. These results demonstrate that G. soja alleles can confer durable transgressive resistance to M. incognita . Correlation between nematode reproduction and root weight In H. glycines trials, mean fresh root weight of CSSLs was 2.17 ± 0.21 g (range 1.05-2.92 g). ZYD00006 had significantly lower biomass (1.09 ± 0.23 g) than Suinong14 (2.55 ± 0.10 g), suggesting G. soja alleles influenced root development. FI was weakly correlated with root weight (R² = 0.0036), whereas CGR correlated moderately with root weight (R² = 0.3571) and FI (R² = 0.4873) (Fig. 3a-c). This suggests root size partly affects CGR but not FI, indicating possible root-independent resistance mechanisms. In M. incognita assays, Suinong14 had higher root weights than ZYD00006 at both inoculum levels. CSSL root weight decreased at the higher inoculum dose, suggesting nematode pressure can suppress root development. Larger root systems tended to have lower EMR and EGR, but most reproduction traits (GI, RF, EMR, EGR) showed weak correlations (R² 0.55) but weak links for GI across inoculum levels (Fig. S2). Cross-dose trait correlations were highest for RF-EGR (R² = 0.8956 at 5,000 eggs; R² = 0.7426 at 10,000 eggs) (Fig. 4), highlighting these as key linked indicators of M. incognita reproduction. In summary, root biomass moderately influenced CGR under SCN infection but had limited, dose-dependent effects under RKN pressure. CSSLs combining robust root systems with low nematode reproduction represent promising sources of resistance. QTL mapping for resistance of H. glycines and M. incongnita For H. glycines (SCN race 4), 25 FI- and 25 CGR-associated loci were identified, with six FI-QTLs and nine CGR-QTLs surpassing P < 0.005 (Table S3). Major FI-QTLs were on Chr5, 7, 8, 9, and 20; major CGR-QTLs were on Chr1, 2, 3, 11, and 12 (Table 1, Fig. 5). No overlap was observed between major FI and CGR loci, indicating distinct genetic control. The phenotypic variance explained (PVE) ranged from 4.8% to 14.3%, with additive effects from -20.14 to 11.37. The strongest FI-QTL ( Hg4-Gs09a-FI ) originated from ZYD00006, whereas the most significant CGR-QTL ( Hg4-Gm02-CGR ) was from Suinong14. For M. incognita , 17 significant QTLs were mapped under 5,000 eggs and 25 under 10,000 eggs, with 18 classified as major QTLs (Table S3, Fig. 5). Many were inoculum dose-sensitive, with PVE ranging from 5.1% to 14.1% (Table 1). QTL hotspots were identified on Chr5, 6, 16, and 19, often harboring co-localized loci for multiple traits (e.g., Mi-Gm05b-RF/EGR , Mi-Gm16a-RF/EMR ) (Fig. 6). Resistance alleles originated from both parents, demonstrating transgressive segregation. Common QTLs for H. glycines (races 4, 5) and M. incognita Comparison with previous work on SCN race 5 (Huang et al. 2021) revealed limited overlap in mapped QTLs among SCN races and RKN (Table 2). Only one CGR-QTL (PVE = 14.3%) was common to SCN race 4 in this study, compared with seven common QTLs for FI/CGR in race 5. Seven major intervals were co-mapped for resistance to SCN races 4 and 5, mainly on Chr2, Chr5, and Chr7. For M. incognita , 18 major QTLs and 139 minor QTLs were identified, with two major intervals shared between H. glycines and M. incognita resistance. These were located on Chr5 (33.50-39.36 Mb) and Chr7 (0.96-2.50 Mb). Overlapping intervals suggest partial sharing of resistance mechanisms across nematode species, but also highlight the complexity of soybean’s polygenic resistance. Discussion Resistance to plant-parasitic nematodes is most effectively dissected when multiple nematode species are evaluated within the same host genetic background. In this study, we extended previous work on H. glycines (HG) type 2.5.7 (race 5) using the Suinong14 × ZYD00006 CSSL population (Huang et al. 2021) to include HG type 1.2.3.5.6.7 (race 4) and M. incognita race 1. This allowed for direct comparisons of resistance QTL across species and pathotypes, revealing both shared and unique genetic determinants. Phenotypic patterns highlighted clear race-specific differences. H. glycines race 4 displayed higher pathogenicity than race 5, producing higher FI values in both parental lines and generally lower resistance in CSSLs. Nevertheless, transgressive segregation for both FI and CGR was detected, indicating that introgressed G. soja alleles can significantly modify host resistance. Correlation analyses revealed that race 4 resistance was more strongly associated with FI-CGR relationships, whereas race 5 resistance was more influenced by CGR-root weight associations. These patterns suggest race-dependent activation of resistance pathways, potentially reflecting distinct transcriptomic or post-transcriptional mechanisms (Huang et al. 2022b). Five FI-QTLs and five CGR-QTLs for race 4 overlapped with loci reported in previous biparental and GWAS studies (www.soybase.org, Table 3), confirming their relevance across pathotypes. Notably, novel loci were identified on Chr9 ( Hg4-Gs09-FIa ) and Chr20 ( Hg4-Gs20-FI ). Among the five FI-QTLs, most resistance alleles originated from G. soja , except for Hg4-Gm07-FIa , which conferred a positive additive effect on resistance. Interestingly, several G. soja -derived FI-QTLs displayed negative additive effects, suggesting possible trade-offs or context-dependent impacts on resistance. The absence of rhg1 - and Rhg4 -associated QTLs aligns with the susceptible genotype profile ( rhg1c + Rhg4b ) of both parents (Huang et al. 2022a). The QTL landscape for M. incognita differed markedly from SCN. While previous studies have consistently reported a major Chr10 QTL (Li et al. 2018; Vuong et al. 2021; Canella et al. 2021, 2022), we did not detect this locus, instead identifying minor QTLs on Chr10 and a key multi-trait locus on Chr18 ( MI-Gs18b-GI ) associated with GI, RF, EMR, and EGR. Additional novel major QTLs for EMR and EGR were mapped to Chr7 and Chr20, supporting the polygenic nature of RKN resistance. Evaluating multiple resistance indices proved essential: GI correlations with other traits were generally weak, while EMR and EGR often aligned with reproduction-based resistance (Fourie et al. 2008). Resistance to both SCN and RKN showed extensive transgressive segregation. For M. incognita , over 90% of CSSLs exhibited enhanced EGR resistance at 5,000 eggs per plant, and 43% at 10,000 eggs per plant. These results echo earlier reports in cotton and soybean where medium-effect alleles combine to generate strong resistance via gene clusters, duplicated loci, or regulatory elements (Wang et al. 2008, 2012, 2017, 2020; Huang et al. 2021). Comparable transgressive effects have been documented in other host-pathogen systems such as tomato-early blight (Anderson et al. 2021) and strawberry- Macrophomina phaseolina (Knapp et al. 2024). Genomic regions conferring broad-spectrum resistance across nematode species are of particular interest for breeding. In this study, QTL co-localizations were found for RF, EMR, and EGR on Chr5, Chr6, Chr16, and Chr19, and two QTL intervals (on Chr5 and Chr7) were shared among all three nematode challenges. Such conserved regions may harbor durable resistance genes suitable for marker-assisted selection. Functional genomic advances have already revealed key resistance genes such as GmSNAP02 (Usovsky et al. 2023), GmMYB29 (Jiang et al. 2025), GmTGA10 , and GmSCT-10 (Lakhssassi et al. 2025) for SCN, and XTH43 (Niraula et al. 2020), GmPR10 (Arraes et al. 2022), and GmEXPA1 (Basso et al. 2023) for RKN. Integrating such knowledge with QTL mapping data accelerates the identification of candidate genes underlying these loci. The use of wild soybean germplasm, as demonstrated here with ZYD00006, remains a powerful approach to broaden the genetic base of resistance in cultivated soybean. Our findings underscore that resistance to both SCN and RKN is quantitatively inherited, polygenic, and strongly influenced by race- or dose-specific interactions. By combining multi-trait phenotyping with high-resolution CSSL mapping, we identified novel loci, transgressive resistance patterns, and shared genomic regions that together provide valuable resources for marker-assisted breeding and functional studies aimed at durable nematode resistance in soybean. Declarations Author contributions Minghui Huang and Chunyan Liu: Data curation; Formal analysis; Investigation; Methodology; Validation; Visualization; Writing original draft. Lei Zhao: Data curation; Investigation. Yanan Zhao: Data curation; Methodology; Resources. Ye Jiang: Data curation; Resources. Doudou Chang: Investigation. Congli Wang : Conceptualization; Data curation; Formal analysis; Funding acquisition; Project administration; Resources; Supervision; Validation; Writing-review and editing. Qingshan Chen: Conceptualization; Data curation; Funding acquisition; Project administration; Resources; Supervision; Writing-review and editing. Chunjie Li: Conceptualization; Data curation; Formal analysis; Funding acquisition; Project administration; Resources; Supervision; Validation; Writing-review and editing. Funding This study was supported by the National Natural Science Foundation of China (31772139, 32272072) to CW, Heilongjiang Province Seed Industry Innovation and Development Project (2024HZYCXNK01), and by National Key Research and Development Program of China (2023YFD1400400). Conflict of interest The authors declare that they have no conflict of interest. Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/. Publisher’s Note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. Supporting information Additional supporting information may be found online in the Supporting Information section at the end of the article. References Acharya K, Tande C, Byamukama E (2016) Determination of Heterodera glycines virulence phenotypes occurring in South Dakota. 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ZYD SN LOD f PVE (%) Additive effect LOD PVE (%) Additive effect QTL-FI Hg4-Gs05-FI 05 29.30-39.42 2726-2892 7.724 *** 93.48 106.58 2.29 5.2 -7.85 Hg4-Gm07a-FI 07a 0.00-7.60 3413-3531 6.691 *** 123.85 104.27 2.2 5.0 9.50 Hg4-Gs08a-FI 08a 9.96-14.77 4234-4471 12.37 ****** 83.31 105.90 2.14 4.8 -20.14 Hg4-Gs09a-FI 09a 0.65-5.47 4833-4865 11.833 ***** 85.78 106.37 2.44 5.5 -10.29 Hg4-Gs20-FI 20 0.48-2.51 10409-10447 5.731 ** 87.26 106.34 2.14 4.8 -11.11 QTL-CGR Hg4-Gs01b-CGR 01b 33.74-49.34 423-482 11.151 ***** 53.05 65.50 2.51 5.7 -6.66 Hg4-Gm02-CGR 02 0.60-9.79 563-883 25.949 ******* 83.13 62.26 6.66 14.3 11.37 Hg4-Gs03b-CGR 03b 36.91-45.51 1878-2080 11.847 ***** 77.15 62.68 3.02 6.7 7.01 Hg4-Gm11a-CGR 11a 0.02-10.12 5465-5804 14.522 ****** 75.34 62.42 2.99 6.7 6.30 Hg4-Gm12-CGR 12 3.91-12.82 6141-6256 13.226 ****** 76.70 62.89 2.73 6.1 7.03 QTL-GI 2.16 5.5 0.11 Mi1-Gm01a-GI 01a 2.21-7.36 278-393 10.267 **** 3.31 3.1 2.25 5.7 -0.12 Mi1-Gs07a-GI 07a 3.90-24.25 3455-3925 9.756 **** 2.89 3.17 2.62 6.7 -0.15 Mi1-Gs13b-GI 13b 28.37-42.52 6863-7097 10.217 **** 2.86 3.16 4.043 ** 3.21 3.57 2.22 5.7 -0.15 Mi1-Gs17-GI 17 13.12-39.60 8857-9095 8.816 **** 2.85 3.16 4.995 ** 3.28 3.58 2.17 5.6 -0.22 Mi1-Gs18b-GI 18b 48.54-57.75 9401-9567 4.661 ** 2.95 3.15 11.291 ***** 3.15 3.59 2.12 5.4 0.29 Mi1-Gm19b-GI 19b 30.04-46.61 9696-10228 8.696 **** 3.4 3.13 13.577 ****** 4.02 3.52 2.16 5.5 0.11 QTL-RF Mi-Gm05b-RF 05b 3.66-39.36 2523-2891 27.759 **** 0.82 0.56 2.3 5.9 0.13 Mi-Gm06c-RF 06c 49.54-51.13 3366-3412 4.65 ** 0.71 0.63 2.21 5.9 0.22 Mi-Gs07a-RF 07a 0.96-4.45 3432-3460 4.175 ** 0.40 0.64 2.0 5.1 0.31 Mi-Gm07c-RF 07c 9.62-17.90 3696-3904 9.522 **** 1.09 0.62 2.26 5.8 0.24 Mi-Gm10-RF 10 5.22-5.46 5149-5152 6.318 ** 0.35 0.15 2.21 5.6 0.10 Mi-Gm16a-RF 16a 3.04-33.12 8380-8590 6.837 *** 0.25 0.14 4.31 10.7 0.07 QTL-EMR Mi-Gm06b-EMR 06b 49.70-51.13 3369-3412 6.139 ** 23.49 18.61 2.05 5.3 4.45 Mi-Gm16-EMR 16 3.04-37.10 8380-8683 10.88 ***** 3.89 2.30 4.85 12.1 1.26 Mi-Gm19a-EMR 19a 6.00-10.12 9639-9688 4.734 ** 33.13 18.56 2.75 11 10.44 QTL-EGR Mi-Gm05b-EGR 05b 3.66-39.36 2523-2891 14.273 ****** 4810.44 3640.31 2.35 6.0 542.74 Mi1-Gm16b-EGR 16b 3.04-33.13 8380-8590 9.006 **** 589.61 307.03 5.75 14.1 177.03 Mi-Gm19-EGR 19 3.63-44.50 9635-10135 7.643 *** 5469.18 3902.07 2.37 10.7 2144.58 a QTL-FI or GI: QTLs related to female index (FI) or gall index (GI) phenotype. Hg4-Gs01-FI or Mi1-M01-RF. The name of QTL for FI or GI on Chr01 from Glycine soja (Gs) ZYD00006 to Heterodera glycines race 4 (HG Type 1.2.3.5.6.7) or Meloidogyne incongnita race 1. Similarly, Hg4-Gm01-FI or Mi1-Gm01-RF represents the name of QTL for FI or GI on Chr01 from G. max cv. Suinong14 to H. glycines race 4 or M.incongnita race 1. b Chr: chromosome. c QTL region (Mbp) was from the first to the end physical position (SNP block). d Signif.: The significant P-values: P < 0.05 (**), 0.01 (***), 0.005 (****), 0.001 (*****), 0.0005 (******). K* score of P < 0.05 (**) was shown in the table. e The values of ZYD and SN show mean values of phenotype (FI or GI) related to ZYD00006 alleles or Suinong14 alleles. f LOD scores and the percentage of phenotypic variance explained (PVE) are by means of MQM mapping analysis and the highest value of LOD and PVE values were shown in the SNP block in the QTL region. LOD (Logarithm of odds) > 2 was shown in the table. The additive effect with positive or negative values means increase or decrease in phenotypic values after the introgression segment of ZYD00006 alleles into Suinong14 background. Table 2. Common QTL intervals associated with resistance to Heterodera glycines (HG) and Meloidogyne incognita (MI) in a soybean chromosome segment substitution line (CSSL) population. Traits include female index (FI) and cysts per gram root (CGR) for SCN HG types 2.5.7 (race 5) and 1.2.3.5.6.7 (race 4), and galling index (GI), reproduction factor (RF), egg masses per gram root (EMR), and eggs per gram root (EGR) for MI race 1 Chr (LG) a QTL interval b Mbp SNP block QTL name MI SCN4 SCN5 2 (D1b) 8.80-9.79 854-883 Hg4-Gm02-CGR Hg5-Gs02a-CGR Hg5-Gs02-FI 5 (A1) 29.30-33.50 2726-2799 Mi1-Gm05b-RF Mi1-Gm05b-EGR Hg4-Gs05-FI 33.50-38.50 2799-2885 Mi1-Gm05b-RF Mi1-Gm05b-EGR Hg4-Gs05-FI Hg5-Gs05-FI 38.50-39.36 2885-2891 Mi1-Gm05b-RF Mi1-Gm05b-EGR Hg4-Gs05-FI Hg5-Gs05-FI Hg5-Gm05-CGR 39.36-39.42 2891-2892 Hg4-Gs05-FI Hg5-Gs05-FI Hg5-Gm05-CGR 7 (M) 0.15-0.67 3414-3422 Hg4-Gm07a-FI Hg5-Gm07-CGR 0.67-0.96 3422-3432 Hg4-Gm07a-FI Hg5-Gm07-CGR Hg5-Gs07b-FI 0.96-2.50 3432-3441 Mi1-Gs07a-RF Hg4-Gm07a-FI Hg5-Gm07-CGR Hg5-Gs07b-FI 2.50-4.45 3441-3460 Mi1-Gs07a-RF Hg4-Gm07a-FI 10.81-15.50 3758-3788 Mi1-Gm07c-RF Hg5-Gs07a-FI 16 (J) 32.40-33.13 8552-8590 Mi1-Gm16a-RF Mi1-Gm16b-EGR Mi1-Gm16-EMR Hg5-Gs16-CGR 33.13-36.50 8590-8666 Mi1-Gm16-EMR Hg5-Gs16-CGR a Chr and LG was short for chromosome and linkage group. b The major QTL region ( P 2) were shown in the table. Table 3 The reported QTLs associated with resistance to Heterodera glycines and Meloidogyne incognita were mapped in chromosome segment substitution lines (CSSLs) QTL name a Reported QTL (soybase.org) QTL-FI Hg4-Gs05-FI SCN 18-1 Hg4-Gm07a-FI SCN 46-1; SCN5-g16 Hg4-Gs08a-FI SCN 8-4; SCN 9-1; SCN 9-3; SCN 29-5; SCN 30-3; SCN 39-8; SCN 42-1; SCN 45-4; SCN 50-1 Hg4-Gs09a-FI SCN5-g19 Hg4-Gs20-FI SCN5-g52 QTL-CGR Hg4-Gs01b-CGR SCN 39-6; SCN 39-11; SCN5-g2; SCN4-g1 Hg4-Gm02-CGR SCN5-g4; SCN5-g3 Hg4-Gs03b-CGR SCN 33-7; SCN 44-15; SCN 51-1; SCN5-g6; SCN1-g1; SCN5-g5 Hg4-Gm11a-CGR SCN 46-6; SCN 49-1; SCN5-g22; SCN5-g25 Hg4-Gm12-CGR SCN 10-4; SCN 10-5; SCN 11-1; SCN 11-2; SCN 39-4; SCN5-g27; SCN5-g26; SCN4-g9 QTL-GI Mi1-Gs18b-GI Mi 1-3; Mi1-4; Mi 2-2; Mi 3-2; Mi 4-1 QTL-RF Mi-Gm10-RF Mi 1-1 a QTL-FI or GI: QTLs related to female index (FI) or gall index (GI) phenotype. Hg4-Gs01-FI or Mi1-M01-RF. The name of QTL for FI or GI on Chr01 from Glycine soja (Gs) ZYD00006 to Heterodera glycines race 4 (HG Type 1.2.3.5.6.7) or Meloidogyne incongnita race 1. Similarly, Hg4-Gm01-FI or Mi1-Gm01-RF represents the name of QTL for FI or GI on Chr01 from G. max cv. Suinong14 to H. glycines race 4 or M.incongnita race 1. Additional Declarations No competing interests reported. Supplementary Files Supplementarytables.xlsx FigureS1.tif FigureS2.tif Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-7587031","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":525812623,"identity":"bbaff0e8-3614-467c-bfd9-6501dc08d587","order_by":0,"name":"Minghui Huang","email":"","orcid":"","institution":"Northeast Institute of Geography and Agroecology, Chinese Academy of Sciences","correspondingAuthor":false,"prefix":"","firstName":"Minghui","middleName":"","lastName":"Huang","suffix":""},{"id":525812624,"identity":"d96be2a5-5f78-4c88-9b7b-1f46bed96257","order_by":1,"name":"Chunyan Liu","email":"","orcid":"","institution":"Northeast 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1","display":"","copyAsset":false,"role":"figure","size":13831279,"visible":true,"origin":"","legend":"\u003cp\u003ePhenotypic distribution of female index (FI) and cysts per gram root (CGR) for \u003cem\u003eHeterodera glycines\u003c/em\u003e HG type 1.2.3.5.6.7 (race 4) in 199 CSSLs derived from the cross between \u003cem\u003eGlycine max\u003c/em\u003e Suinong14 and \u003cem\u003eG. soja\u003c/em\u003e ZYD00006\u003c/p\u003e","description":"","filename":"Figure1.png","url":"https://assets-eu.researchsquare.com/files/rs-7587031/v1/de48c93560b0dbd3a4d9395a.png"},{"id":93142812,"identity":"a3fd09f6-8f9a-4527-aa7d-401c54d57074","added_by":"auto","created_at":"2025-10-09 13:16:00","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":22501201,"visible":true,"origin":"","legend":"\u003cp\u003ePhenotypic performance of 177 CSSLs in response to two inoculum levels (Test1-5,000 and Test2-10,000 eggs per plant) of \u003cem\u003eMeloidogyne incognita\u003c/em\u003e. Traits include gall index (GI), reproduction factor (RF), egg masses per gram root (EMR), and eggs per gram root (EGR)\u003c/p\u003e","description":"","filename":"Figure2.png","url":"https://assets-eu.researchsquare.com/files/rs-7587031/v1/3adc326e6f5d7a67a14f75ab.png"},{"id":93141198,"identity":"6be92168-7a07-4097-a181-7ce9868543e4","added_by":"auto","created_at":"2025-10-09 13:08:00","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":14697908,"visible":true,"origin":"","legend":"\u003cp\u003eCorrelation between female index (FI), cysts per gram root (CGR), and fresh root weight under \u003cem\u003eHeterodera glycines\u003c/em\u003e infection in 199 CSSLs derived from \u003cem\u003eGlycine max\u003c/em\u003e Suinong14 and \u003cem\u003eG. soja\u003c/em\u003e ZYD00006\u003c/p\u003e","description":"","filename":"Figure3.png","url":"https://assets-eu.researchsquare.com/files/rs-7587031/v1/9b5668910a782e475b449f34.png"},{"id":93141200,"identity":"b6734ecf-2ece-412f-a07d-bbc570e92b2b","added_by":"auto","created_at":"2025-10-09 13:08:00","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":19148327,"visible":true,"origin":"","legend":"\u003cp\u003eCorrelation between reproduction traits (RF, EMR, EGR) at two inoculum levels (Test1-5,000 and Test2-10,000 eggs per plant) of \u003cem\u003eMeloidogyne incognita\u003c/em\u003e in 177 CSSLs\u003c/p\u003e","description":"","filename":"Figure4.png","url":"https://assets-eu.researchsquare.com/files/rs-7587031/v1/85ea05f66e0addf8a02a0cdb.png"},{"id":93144069,"identity":"e396e6ef-ba8a-4364-b1b7-b70422f51e6f","added_by":"auto","created_at":"2025-10-09 13:24:01","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":63750489,"visible":true,"origin":"","legend":"\u003cp\u003eChromosomal distribution of significant QTLs for \u003cem\u003eHeterodera glycines\u003c/em\u003e and \u003cem\u003eMeloidogyne incognita\u003c/em\u003e across all linkage groups in soybean CSSLs. MI-1 and MI-2 represent the two inoculum levels\u003c/p\u003e","description":"","filename":"Figure5.png","url":"https://assets-eu.researchsquare.com/files/rs-7587031/v1/5160d44364e02beeacbc672b.png"},{"id":93142817,"identity":"c97597cb-af43-4075-a974-d97c91b533e9","added_by":"auto","created_at":"2025-10-09 13:16:01","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":36025436,"visible":true,"origin":"","legend":"\u003cp\u003ePhenotypic variance explained by major QTLs located in shared chromosomal intervals associated with \u003cem\u003eMeloidogyne incognita\u003c/em\u003e resistance traits\u003c/p\u003e","description":"","filename":"Figure6.png","url":"https://assets-eu.researchsquare.com/files/rs-7587031/v1/cf0e37dccd135c5b66580073.png"},{"id":93141201,"identity":"720ff3b4-be07-4818-af66-e5b9cc6e832c","added_by":"auto","created_at":"2025-10-09 13:08:00","extension":"xlsx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":2476234,"visible":true,"origin":"","legend":"","description":"","filename":"Supplementarytables.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-7587031/v1/11c7a75e9b2c494dac3c1bbe.xlsx"},{"id":93142813,"identity":"906e27ce-af79-46d2-86d9-22782300045a","added_by":"auto","created_at":"2025-10-09 13:16:00","extension":"tif","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":5561612,"visible":true,"origin":"","legend":"","description":"","filename":"FigureS1.tif","url":"https://assets-eu.researchsquare.com/files/rs-7587031/v1/abb40cf6090df5c5b98363e1.tif"},{"id":93141212,"identity":"5f6bc4c0-c963-4588-8d42-6ccc5462391a","added_by":"auto","created_at":"2025-10-09 13:08:01","extension":"tif","order_by":3,"title":"","display":"","copyAsset":false,"role":"supplement","size":5424130,"visible":true,"origin":"","legend":"","description":"","filename":"FigureS2.tif","url":"https://assets-eu.researchsquare.com/files/rs-7587031/v1/dafc0720ec23d6794eda5d28.tif"}],"financialInterests":"No competing interests reported.","formattedTitle":"Transgressive inheritance of broad-spectrum resistance to Heterodera glycines and Meloidogyne incognita in soybean chromosome segment substitution lines","fulltext":[{"header":"Introduction","content":"\u003cp\u003eSoybean (\u003cem\u003eGlycine max\u003c/em\u003e L. Merr.) is a globally important oilseed crop and a key component of food and agricultural systems. China ranks as the fourth-largest soybean producer, following the United States, Brazil, and Argentina. Among the major biotic constraints, plant-parasitic nematodes (PPNs) are the second most damaging group of pathogens globally, significantly threatening food security and crop yields. Among PPNs, the soybean cyst nematode (\u003cem\u003eHeterodera glycines\u003c/em\u003e, SCN) and root-knot nematodes (\u003cem\u003eMeloidogyne\u003c/em\u003e spp., RKN) are the most prevalent and destructive nematodes, causing serious yield losses across diverse cropping systems (Bent et al. 2022; Thakur et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Both SCN and RKN are obligate sedentary endoparasites. Their infective second-stage juveniles (J2s) invade roots and secrete effectors through their stylets to establish specialized feeding sites \u0026mdash; syncytia in SCN and giant cells in RKN \u0026mdash; disrupting nutrient flow and reducing yield (Niblack et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Mitchum and Baum \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2008\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eIn China, SCN is especially problematic in the Northeast, the country\u0026rsquo;s primary soybean-producing region. Yield losses typically range from 20\u0026ndash;30%, rising to 60\u0026ndash;70% under severe infestations (Peng 2021). Annual losses due to SCN exceed \u003cspan\u003e$\u003c/span\u003e1\u0026nbsp;billion in the U.S. (Koenning and Wrather \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2006\u003c/span\u003e) and 600\u0026nbsp;million yuan in China (Peng 2021). RKNs, particularly \u003cem\u003eMeloidogyne incognita\u003c/em\u003e, infect over 5,500 plant species and cause global crop losses of approximately \u003cspan\u003e$\u003c/span\u003e7.8\u0026nbsp;billion (Caillaud et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Nicol et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). In China, RKN-induced yield reductions in vegetables can exceed 30%, with annual economic losses surpassing 3\u0026nbsp;billion yuan (Peng 2021). \u003cem\u003eM. incognita\u003c/em\u003e is the dominant RKN species worldwide and in China. Although historically confined to central regions, it is now spreading to northern provinces via protected cultivation systems. Greenhouse conditions (~\u0026thinsp;23\u0026deg;C) promote overwintering and rapid reproduction (Zhao et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Hong et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), accelerating its spread in regions such as Liaoning and Heilongjiang and increasing its threat to soybean production.\u003c/p\u003e\u003cp\u003eCurrent control strategies for SCN and RKN rely primarily on resistant cultivars and crop rotation. However, both \u003cem\u003eH. glycines\u003c/em\u003e and \u003cem\u003eM. incognita\u003c/em\u003e exhibit high genetic diversity and complex race structures (HG types and pathotypes), making breeding efforts challenging. Most commercial SCN-resistant cultivars are derived from only two sources (Peking and PI 88788), leading to the emergence of virulent nematode populations, particularly in Heilongjiang, where dominant SCN races have shifted from race 3 to more aggressive races 4 and 14 (Acharya et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Bayless et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Huang et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2022a\u003c/span\u003e; You et al. \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Although \u003cem\u003eM. incognita\u003c/em\u003e has been considered a warm-climate pest, its rapid expansion in protected environments highlights the urgency of addressing its threat in colder regions, including Northeast China (Zhao et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Li et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). While international studies dominate RKN resistance research (Schmitt and Bell\u0026eacute; 2016; Canella et al. 2022; Gontijio et al. 2024), domestic progress is increasing. For example, Li et al. (\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) screened native soybean germplasm under greenhouse conditions in Heilongjiang for \u003cem\u003eM. incognita\u003c/em\u003e resistance. SCN resistance is typically assessed using the Female Index (FI), while the cysts per gram of root (CGR) provides a root-weight-adjusted metric (Huang et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). In contrast, no standardized system exists for \u003cem\u003eM. incognita\u003c/em\u003e resistance evaluation. Most studies use the Gall Index (GI), often without accounting for egg counts or standardized inoculum, which hampers precision. Thus, developing a robust and comprehensive evaluation framework for RKN resistance remains a key priority.\u003c/p\u003e\u003cp\u003eResistance to both SCN and RKN in soybean is polygenic and quantitatively inherited (Concibido et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2004\u003c/span\u003e;Li et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Vieira et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Bent et al. 2022). To date, over 300 quantitative trait loci (QTLs) associated with SCN resistance have been mapped across all 20 soybean chromosomes (SoyBase, \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e\u003ca href=\"http://www.soybase.org\" target=\"_blank\"\u003ewww.soybase.org\u003c/a\u003e\u003c/span\u003e\u003cspan address=\"http://www.soybase.org\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e), reflecting the complex genetic architecture of resistance. Among them, two major loci \u0026mdash; \u003cem\u003erhg1\u003c/em\u003e on chromosome 18 and \u003cem\u003eRhg4\u003c/em\u003e on chromosome 8, have been most extensively characterized (Cook et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2012\u003c/span\u003e, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Liu et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2012\u003c/span\u003e, \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). The \u003cem\u003erhg1\u003c/em\u003e locus comprises multiple genes, including α-SNAP, amino acid transporter (AAT), and WI12, as well as other SNAP family members such as \u003cem\u003eGmSNAP11\u003c/em\u003e and \u003cem\u003eGmSNAP02\u003c/em\u003e, which play pivotal roles in nematode resistance (Bayless et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2016\u003c/span\u003e, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Guo et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Dong et al. 2022; Han et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Usovsky et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Du et al. 2024). In contrast, \u003cem\u003eRhg4\u003c/em\u003e encodes a serine hydroxymethyltransferase (\u003cem\u003eGmSHMT08\u003c/em\u003e), and its resistance effect is influenced by copy number variation and allelic dosage at both \u003cem\u003erhg1\u003c/em\u003e and \u003cem\u003eRhg4\u003c/em\u003e loci (Patil et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Despite this progress, additional minor and environment-sensitive QTLs continue to be discovered, underscoring the need for deeper exploration of resistance loci, particularly in wild-derived populations and under field-relevant pathogen pressures. However, resistance QTLs for \u003cem\u003eM. incognita\u003c/em\u003e are less well defined. Early mapping in PI 96354 identified two QTLs on Chr10 and Chr18 (Tamulonis et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e1997\u003c/span\u003e), with later studies reporting additional loci on Chr6, 7, 8, 13, and 17 (Ha et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Fourie et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Xu et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Jiao et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Vuong et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Li et al. (\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) reported that four major QTLs (Chr 10, 18, 20, and 7) associated with \u003cem\u003eM. incognita\u003c/em\u003e resistance. A recent genome-wide association study (GWAS) involving 717 accessions detected major associations on Chr10 and minor loci on Chr11 (Vieira et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). However, many of these intervals are large and based solely on GI, limiting their practical utility.\u003c/p\u003e\u003cp\u003eTo improve QTL resolution, high-density genetic maps and multidimensional resistance indices are essential. Despite identification of major resistance loci, novel and durable sources of resistance are urgently required to enhance breeding pipelines. Traditional QTL mapping populations (e.g., F\u003csub\u003e₂\u003c/sub\u003e, RILs, DH) often suffer from broad confidence intervals due to genetic background noise. Chromosome Segment Substitution Lines (CSSLs), developed by repeated backcrossing and selfing, enable precise QTL mapping in near-isogenic backgrounds. By substituting specific genomic segments from a donor parent into a recurrent background, CSSLs improve the detection of both major and minor QTLs (Balakrishnan et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). The CSSL population used in this study, derived from Suinong14 (recurrent parent) and ZYD00006 (wild soybean donor), has been under development in Northeast China for over a decade (Xin et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Jiang et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Wang et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Jong et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). This population has previously been applied to map QTLs for seed quality, nodulation, drought, cold tolerance, and more recently, transgressive resistance to \u003cem\u003eH. glycines\u003c/em\u003e HG Type 2.5.7 (race 5), based on FI and CGR (Huang et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). However, that study did not address virulent \u003cem\u003eH. glycines\u003c/em\u003e race 4 or the root-knot nematode \u003cem\u003eM. incognita\u003c/em\u003e, which are emerging threats to soybean production and require further investigation.\u003c/p\u003e\u003cp\u003eIn this study, we aimed to (1) assess the resistance of the Suinong14 \u0026times; ZYD00006 CSSL population to \u003cem\u003eH. glycines\u003c/em\u003e race 4 and \u003cem\u003eM. incognita\u003c/em\u003e race 1 under different inoculation pressures, (2) identify novel QTLs associated with resistance traits, and (3) evaluate transgressive segregation patterns and potential elite lines for breeding. Resistance was evaluated using multiple indices: FI and CGR for soybean cyst nematode, and GI, reproduction factor (RF), egg masses per gram root (EMR), and eggs per gram root (EGR) for root-knot nematode. High-resolution QTL mapping was conducted using a published physical map (Huang et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). These efforts will deepen our understanding of the genetic architecture of nematode resistance and support the development of soybean cultivars with durable, broad-spectrum resistance suited to the agroecological conditions of Northeast China.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cp\u003e\u003cstrong\u003ePlant and nematode materials\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eSoybean chromosome segment substitution lines (CSSLs) at the BC\u003csub\u003e3\u003c/sub\u003eF\u003csub\u003e7\u003c/sub\u003e-BC\u003csub\u003e8\u003c/sub\u003eF\u003csub\u003e2\u003c/sub\u003e generations were developed through backcrossing between the recurrent parent \u003cem\u003eG. max\u003c/em\u003e \u0026lsquo;Suinong14\u0026rsquo; and the wild donor parent \u003cem\u003eG. soja\u003c/em\u003e \u0026lsquo;ZYD00006\u0026rsquo; (Huang et al. 2021; Li et al. 2021; Pan et al. 2022; Wang et al. 2023; Jong et al. 2024). The CSSL population, along with both parental lines, was obtained from Northeast Agricultural University. For resistance screening, 199 CSSLs were evaluated against \u003cem\u003eH. glycines\u003c/em\u003e HG type 1.2.3.5.6.7 (designated as SCN race 4), and 177 CSSLs were evaluated against \u003cem\u003eM. incognita\u003c/em\u003e (RKN race 1). Susceptible soybean cultivars \u0026lsquo;Dongsheng 1\u0026rsquo; and \u0026lsquo;Lee68\u0026rsquo; were used as propagation hosts and resistance controls for SCN, respectively, while \u003cem\u003eM. incognita\u003c/em\u003e was maintained on tomato (\u003cem\u003eSolanum lycopersicum\u003c/em\u003e) cultivar \u0026lsquo;Zhongshu 4\u0026rsquo;. Nematode cultures were maintained in a greenhouse under controlled conditions (22-28 \u0026deg;C, 16 h light/8 h dark photoperiod) at the Northeast Institute of Geography and Agroecology, Chinese Academy of Sciences.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003ePreparation of SCN and RKN inocula\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eSCN cysts were extracted from infected soybean roots and surrounding soil 35-40 days post-inoculation. Cysts were crushed to release eggs, and second-stage juveniles (J2s) were hatched in sterile water containing 3 mM ZnSO₄ at 28 \u0026deg;C. For RKN, roots were harvested 42 days post-inoculation, washed thoroughly, and immersed in 50 mL of 1% sodium hypochlorite (NaClO) solution, then shaken either on a shaker or by hand for 3 minutes. The suspension was filtered through a nested sieve set (75 \u0026mu;m upper, 25 \u0026mu;m lower) and rinsed with pressurized water. Eggs retained on the lower sieve were transferred for hatching as described for SCN. J2s and eggs were counted under a stereo microscope (Olympus, Japan) for subsequent inoculation.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eResistance evaluation of soybean to SCN and RKN\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eSeeds were sown in 8 cm \u0026times; 12 cm pots filled with sterilized soil and sand (1:1 v/v). After emergence, seedlings were thinned to one per pot. At 8-9 days after planting, two 4-5 cm-deep holes were made near each seedling for nematode inoculation. Each plant was inoculated with either 1,000 SCN J2s or 5,000/10,000 RKN eggs. SCN resistance was assessed 21 days post-inoculation by counting cysts on washed roots under fluorescent light. For RKN, roots were harvested 42 days post-inoculation, dried, and stained in 75 mg/L erioglaucine for 30 min. Egg masses were counted under a light microscope, and fresh root weights were recorded. Eggs were extracted as described in Section 2.2. All experiments used a randomized complete block design with 5-6 biological replicates per line and were repeated twice. RKN evaluations were conducted at two inoculation doses, and \u0026lsquo;Lee68\u0026rsquo; served as a susceptible control in all experiments.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eSCN and RKN\u003cem\u003e\u0026nbsp;\u003c/em\u003eresistance evaluation\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eSCN resistance was quantified using the FI, calculated as the mean cyst count on a test line divided by the mean count on the susceptible control \u0026lsquo;Lee68\u0026rsquo;, multiplied by 100. Classification followed Schmitt and Shannon (1992): FI 1-10 = resistant, 11-30 = moderately resistant, 31-60 = moderately susceptible, \u0026gt; 60 = susceptible. An additional index, CGR was calculated by dividing cyst number by root weight to adjust for biomass. For RKN, four indices were used: GI, RF, EMR and EGR. EMR was calculated by dividing egg mass number by root weight, and EGR by dividing egg number by root weight. RF was calculated as total eggs per plant divided by eggs inoculated. Resistance levels based on EGR were: highly resistant (\u0026lt; 1,000), resistant (1,000 - 4,999), susceptible (5,000 - 9,999), and highly susceptible (\u0026ge; 10,000).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eQTL detection for SCN and RKN resistance\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eQuantitative trait loci (QTLs) for SCN (FI and CGR) and RKN (GI, RF, EMR, EGR) resistance were identified using MapQTL 5.0 (Van Ooijen 2004) with phenotypic data and a SNP-based physical map comprising 3,780 bin markers (Huang et al. 2021). The Kruskal-Wallis (K*) test and multiple QTL mapping (MQM) were applied. Significant QTLs were defined by \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.005 in K* or LOD \u0026gt; 2 in MQM. LOD thresholds were determined by 1,000 permutations at \u0026alpha; = 0.05 (Churchill and Doerge 1994). QTLs with \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.1 were visualized using MapChart 2.3 (Voorrips 2002). QTLs were named following Huang et al. (2021). SCN QTLs were denoted as \u003cem\u003eHg4-Gs/GmXXa/b-FI/CGR\u003c/em\u003e, and RKN QTLs as \u003cem\u003eMi1-Gs/GmXXa/b-GI/RF/EM/EGR\u003c/em\u003e. Here, \u0026lsquo;\u003cem\u003eHg\u003c/em\u003e4\u0026rsquo; and \u0026lsquo;\u003cem\u003eMi\u003c/em\u003e1\u0026rsquo; refer to \u003cem\u003eH. glycines\u003c/em\u003e race 4 and \u003cem\u003eM. incognita\u003c/em\u003e race 1, \u0026lsquo;\u003cem\u003eGs\u003c/em\u003e\u0026rsquo; or \u0026lsquo;\u003cem\u003eGm\u003c/em\u003e\u0026rsquo; indicate the allele origin (\u003cem\u003eG. soja\u003c/em\u003e or \u003cem\u003eG. max\u003c/em\u003e), \u0026lsquo;XX\u0026rsquo; the chromosome number, and \u0026lsquo;a/b\u0026rsquo; distinguish multiple QTLs on the same chromosome.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eStatistical analyses\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003ePhenotypic data were analyzed by one-way ANOVA using IBM SPSS Statistics 26.0. Tukey\u0026rsquo;s HSD test (\u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05) was used to compare cyst and FI values among lines. Correlations between resistance parameters were evaluated using Microsoft Excel 2013.\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003e\u003cstrong\u003ePhenotypic identification of CSSLs and p\u003c/strong\u003e\u003cstrong\u003earents to\u0026nbsp;\u003c/strong\u003e\u003cstrong\u003e\u003cem\u003eH. glycines\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;HG type 1.2.3.5.6.7\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003ePhenotypic evaluation of CSSLs and their parental lines against \u003cem\u003eH. glycines\u003c/em\u003e HG type 1.2.3.5.6.7 (SCN race 4) revealed no significant difference in cysts/plant and FI among the parent Suinong14 (127.4 \u0026plusmn; 0.94; 105.11 \u0026plusmn; 9.03), ZYD00006 (108 \u0026plusmn; 9.75; 89.11 \u0026plusmn; 8.05), and the susceptible control Lee68 (122.1 \u0026plusmn; 13.92; 100 \u0026plusmn; 11.48) (Fig. 1, Table S1). However, \u003cem\u003eG. soja\u003c/em\u003e ZYD00006 exhibited a markedly higher number of CGR (113.52 \u0026plusmn; 17.29) compared to \u003cem\u003eG. max\u003c/em\u003e Suinong14 (50.11 \u0026plusmn; 4.75) and the susceptible cultivar Lee68 (43.29 \u0026plusmn; 7.95). Among the 199 CSSLs evaluated, FI values ranged from 45.87 to 164.05, and CGR values ranged from 24.63 to 118.07, indicating considerable phenotypic diversity. Several CSSLs displayed reduced susceptibility compared to both parents. Notably, Lines 205 and 193 showed the lowest FI (45.87 \u0026plusmn; 12.08; 58.17 \u0026plusmn; 33.42) and CGR (36.07 \u0026plusmn; 9.23; 24.63 \u0026plusmn; 13.91), with reductions of 48.5% and 50.9%, respectively, compared with ZYD00006 and Suinong14. Forty-four CSSLs had CGR lower than Suinong14, representing 22.1% transgressive resistance. While no line had FI \u0026lt; 30 (highly resistant threshold), nearly half (49.8%) of CSSLs exhibited FI values lower than Suinong14 (FI = 105.11 \u0026plusmn; 9.03), indicating moderate resistance improvements. Eight CSSLs (Lines 4, 24, 50, 57, 85, 92, 193, and 205) combined CGR \u0026lt; 60 with FI \u0026lt; 70, identifying them as promising candidates for SCN resistance breeding. Some lines displayed incongruent FI and CGR values (e.g., Line 213: CGR = 44.94 \u0026plusmn; 3.84; FI = 103.41 \u0026plusmn; 8.43), suggesting different loci may control these traits. These results indicate that \u003cem\u003eG. soja\u003c/em\u003e introgressions can enhance SCN resistance, even in the absence of highly resistant parents, and that selected CSSLs may serve as valuable pre-breeding resources under high disease pressure.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEvaluation of CSSLs under different inoculum levels of \u003cem\u003eM. incongnita\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003ePhenotypic responses of CSSLs to \u003cem\u003eM. incognita\u003c/em\u003e race 1 at 5,000 and 10,000 eggs per plant are summarized in Fig. 2 and Table S1. No significant differences were observed between Suinong14 and ZYD00006 for GI, RF, EMR, or EGR at either inoculum level. At 5,000 eggs per plant, most CSSLs performed better than Suinong14: 40 lines had reduced GI, 6 had lower RF, 135 had decreased EMR, and 161 had reduced EGR. Many lines were highly resistant, with minimal nematode reproduction. At 10,000 eggs per plant, transgressive resistance remained: 45% of lines had increased GI, 33% reduced RF, 37% reduced EMR, and 43% reduced EGR. However, the proportion of resistant lines (EGR \u0026lt; 5,000) decreased by 21.47% relative to the lower inoculum, indicating dose-dependent declines in resistance. Several CSSLs, including Lines 206, 210, 213, and 214, maintained stable resistance across doses, and eleven lines (e.g., 46, 119, 152, 154, 155, 156, 158, 159, 175, 189, 190) had EGR between 1,000-2,000 at high dose and \u0026lt; 1,000 at low dose. These results demonstrate that \u003cem\u003eG. soja\u003c/em\u003e alleles can confer durable transgressive resistance to \u003cem\u003eM. incognita\u003c/em\u003e.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCorrelation between nematode\u003cem\u003e\u0026nbsp;\u003c/em\u003ereproduction\u003cem\u003e\u0026nbsp;\u003c/em\u003eand root weight\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eIn \u003cem\u003eH. glycines\u003c/em\u003e trials, mean fresh root weight of CSSLs was 2.17 \u0026plusmn; 0.21 g (range 1.05-2.92 g). ZYD00006 had significantly lower biomass (1.09 \u0026plusmn; 0.23 g) than Suinong14 (2.55 \u0026plusmn; 0.10 g), suggesting \u003cem\u003eG. soja\u003c/em\u003e alleles influenced root development. FI was weakly correlated with root weight (R\u0026sup2; = 0.0036), whereas CGR correlated moderately with root weight (R\u0026sup2; = 0.3571) and FI (R\u0026sup2; = 0.4873) (Fig. 3a-c). This suggests root size partly affects CGR but not FI, indicating possible root-independent resistance mechanisms.\u003c/p\u003e\n\u003cp\u003eIn \u003cem\u003eM. incognita\u003c/em\u003e assays, Suinong14 had higher root weights than ZYD00006 at both inoculum levels. CSSL root weight decreased at the higher inoculum dose, suggesting nematode pressure can suppress root development. Larger root systems tended to have lower EMR and EGR, but most reproduction traits (GI, RF, EMR, EGR) showed weak correlations (R\u0026sup2; \u0026lt; 0.1), except RF at high inoculum (R\u0026sup2; = 0.3311) (Fig. S1). Correlation analysis showed strong associations among RF, EMR, and EGR (R\u0026sup2; \u0026gt; 0.55) but weak links for GI across inoculum levels (Fig. S2). Cross-dose trait correlations were highest for RF-EGR (R\u0026sup2; = 0.8956 at 5,000 eggs; R\u0026sup2; = 0.7426 at 10,000 eggs) (Fig. 4), highlighting these as key linked indicators of \u003cem\u003eM. incognita\u003c/em\u003e reproduction.\u003c/p\u003e\n\u003cp\u003eIn summary, root biomass moderately influenced CGR under SCN infection but had limited, dose-dependent effects under RKN pressure. CSSLs combining robust root systems with low nematode reproduction represent promising sources of resistance.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eQTL mapping for resistance of \u003cem\u003eH. glycines\u003c/em\u003e and \u003cem\u003eM. incongnita\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eFor \u003cem\u003eH. glycines\u003c/em\u003e (SCN race 4), 25 FI- and 25 CGR-associated loci were identified, with six FI-QTLs and nine CGR-QTLs surpassing \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.005 (Table S3). Major FI-QTLs were on Chr5, 7, 8, 9, and 20; major CGR-QTLs were on Chr1, 2, 3, 11, and 12 (Table 1, Fig. 5). No overlap was observed between major FI and CGR loci, indicating distinct genetic control. The phenotypic variance explained (PVE) ranged from 4.8% to 14.3%, with additive effects from -20.14 to 11.37. The strongest FI-QTL (\u003cem\u003eHg4-Gs09a-FI\u003c/em\u003e) originated from ZYD00006, whereas the most significant CGR-QTL (\u003cem\u003eHg4-Gm02-CGR\u003c/em\u003e) was from Suinong14.\u003c/p\u003e\n\u003cp\u003eFor \u003cem\u003eM. incognita\u003c/em\u003e, 17 significant QTLs were mapped under 5,000 eggs and 25 under 10,000 eggs, with 18 classified as major QTLs (Table S3, Fig. 5). Many were inoculum dose-sensitive, with PVE ranging from 5.1% to 14.1% (Table 1). QTL hotspots were identified on Chr5, 6, 16, and 19, often harboring co-localized loci for multiple traits (e.g., \u003cem\u003eMi-Gm05b-RF/EGR\u003c/em\u003e, \u003cem\u003eMi-Gm16a-RF/EMR\u003c/em\u003e) (Fig. 6). Resistance alleles originated from both parents, demonstrating transgressive segregation.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCommon QTLs for \u003cem\u003eH. glycines\u003c/em\u003e (races 4, 5) and \u003cem\u003eM. incognita\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eComparison with previous work on SCN race 5 (Huang et al. 2021) revealed limited overlap in mapped QTLs among SCN races and RKN (Table 2). Only one CGR-QTL (PVE = 14.3%) was common to SCN race 4 in this study, compared with seven common QTLs for FI/CGR in race 5. Seven major intervals were co-mapped for resistance to SCN races 4 and 5, mainly on Chr2, Chr5, and Chr7. For \u003cem\u003eM. incognita\u003c/em\u003e, 18 major QTLs and 139 minor QTLs were identified, with two major intervals shared between \u003cem\u003eH. glycines\u003c/em\u003e and \u003cem\u003eM. incognita\u003c/em\u003e resistance. These were located on Chr5 (33.50-39.36 Mb) and Chr7 (0.96-2.50 Mb). Overlapping intervals suggest partial sharing of resistance mechanisms across nematode species, but also highlight the complexity of soybean\u0026rsquo;s polygenic resistance.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eResistance to plant-parasitic nematodes is most effectively dissected when multiple nematode species are evaluated within the same host genetic background. In this study, we extended previous work on \u003cem\u003eH. glycines\u003c/em\u003e (HG) type 2.5.7 (race 5) using the Suinong14 \u0026times; ZYD00006 CSSL population (Huang et al. 2021) to include HG type 1.2.3.5.6.7 (race 4) and \u003cem\u003eM. incognita\u003c/em\u003e race 1. This allowed for direct comparisons of resistance QTL across species and pathotypes, revealing both shared and unique genetic determinants. Phenotypic patterns highlighted clear race-specific differences. \u003cem\u003eH. glycines\u003c/em\u003e race 4 displayed higher pathogenicity than race 5, producing higher FI values in both parental lines and generally lower resistance in CSSLs. Nevertheless, transgressive segregation for both FI and CGR was detected, indicating that introgressed \u003cem\u003eG. soja\u003c/em\u003e alleles can significantly modify host resistance. Correlation analyses revealed that race 4 resistance was more strongly associated with FI-CGR relationships, whereas race 5 resistance was more influenced by CGR-root weight associations. These patterns suggest race-dependent activation of resistance pathways, potentially reflecting distinct transcriptomic or post-transcriptional mechanisms (Huang et al. 2022b).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eFive FI-QTLs and five CGR-QTLs for race 4 overlapped with loci reported in previous biparental and GWAS studies (www.soybase.org, Table 3), confirming their relevance across pathotypes. Notably, novel loci were identified on Chr9 (\u003cem\u003eHg4-Gs09-FIa\u003c/em\u003e) and Chr20 (\u003cem\u003eHg4-Gs20-FI\u003c/em\u003e). Among the five FI-QTLs, most resistance alleles originated from \u003cem\u003eG. soja\u003c/em\u003e, except for \u003cem\u003eHg4-Gm07-FIa\u003c/em\u003e, which conferred a positive additive effect on resistance. Interestingly, several \u003cem\u003eG. soja\u003c/em\u003e-derived FI-QTLs displayed negative additive effects, suggesting possible trade-offs or context-dependent impacts on resistance. The absence of \u003cem\u003erhg1\u003c/em\u003e- and \u003cem\u003eRhg4\u003c/em\u003e-associated QTLs aligns with the susceptible genotype profile (\u003cem\u003erhg1c\u003c/em\u003e + \u003cem\u003eRhg4b\u003c/em\u003e) of both parents (Huang et al. 2022a).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe QTL landscape for \u003cem\u003eM. incognita\u003c/em\u003e differed markedly from SCN. While previous studies have consistently reported a major Chr10 QTL (Li et al. 2018; Vuong et al. 2021; Canella et al. 2021, 2022), we did not detect this locus, instead identifying minor QTLs on Chr10 and a key multi-trait locus on Chr18 (\u003cem\u003eMI-Gs18b-GI\u003c/em\u003e) associated with GI, RF, EMR, and EGR. Additional novel major QTLs for EMR and EGR were mapped to Chr7 and Chr20, supporting the polygenic nature of RKN resistance. Evaluating multiple resistance indices proved essential: GI correlations with other traits were generally weak, while EMR and EGR often aligned with reproduction-based resistance (Fourie et al. 2008).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eResistance to both SCN and RKN showed extensive transgressive segregation. For \u003cem\u003eM. incognita\u003c/em\u003e, over 90% of CSSLs exhibited enhanced EGR resistance at 5,000 eggs per plant, and 43% at 10,000 eggs per plant. These results echo earlier reports in cotton and soybean where medium-effect alleles combine to generate strong resistance via gene clusters, duplicated loci, or regulatory elements (Wang et al. 2008, 2012, 2017, 2020; Huang et al. 2021). Comparable transgressive effects have been documented in other host-pathogen systems such as tomato-early blight (Anderson et al. 2021) and strawberry-\u003cem\u003eMacrophomina phaseolina\u003c/em\u003e (Knapp et al. 2024).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eGenomic regions conferring broad-spectrum resistance across nematode species are of particular interest for breeding. In this study, QTL co-localizations were found for RF, EMR, and EGR on Chr5, Chr6, Chr16, and Chr19, and two QTL intervals (on Chr5 and Chr7) were shared among all three nematode challenges. Such conserved regions may harbor durable resistance genes suitable for marker-assisted selection. Functional genomic advances have already revealed key resistance genes such as \u003cem\u003eGmSNAP02\u003c/em\u003e (Usovsky et al. 2023), \u003cem\u003eGmMYB29\u003c/em\u003e (Jiang et al. 2025), \u003cem\u003eGmTGA10\u003c/em\u003e, and \u003cem\u003eGmSCT-10\u003c/em\u003e (Lakhssassi et al. 2025) for SCN, and \u003cem\u003eXTH43\u003c/em\u003e (Niraula et al. 2020), \u003cem\u003eGmPR10\u003c/em\u003e (Arraes et al. 2022), and \u003cem\u003eGmEXPA1\u003c/em\u003e (Basso et al. 2023) for RKN. Integrating such knowledge with QTL mapping data accelerates the identification of candidate genes underlying these loci. The use of wild soybean germplasm, as demonstrated here with ZYD00006, remains a powerful approach to broaden the genetic base of resistance in cultivated soybean. Our findings underscore that resistance to both SCN and RKN is quantitatively inherited, polygenic, and strongly influenced by race- or dose-specific interactions. By combining multi-trait phenotyping with high-resolution CSSL mapping, we identified novel loci, transgressive resistance patterns, and shared genomic regions that together provide valuable resources for marker-assisted breeding and functional studies aimed at durable nematode resistance in soybean.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAuthor contributions\u0026nbsp;\u003c/strong\u003eMinghui Huang and Chunyan Liu: Data curation; Formal analysis; Investigation; Methodology; Validation; Visualization; Writing original draft. Lei Zhao: Data curation; Investigation. Yanan Zhao: Data curation; Methodology; Resources. Ye Jiang: Data curation; Resources. Doudou Chang: Investigation. Congli Wang : Conceptualization; Data curation; Formal analysis; Funding acquisition; Project administration; Resources; Supervision; Validation; Writing-review and editing. Qingshan Chen: Conceptualization; Data curation; Funding acquisition; Project administration; Resources; Supervision; Writing-review and editing. Chunjie Li: Conceptualization; Data curation; Formal analysis; Funding acquisition; Project administration; Resources; Supervision; Validation; Writing-review and editing.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e This study was supported by the National Natural Science Foundation of China (31772139, 32272072) to CW, Heilongjiang Province Seed Industry Innovation and Development Project (2024HZYCXNK01), and by National Key Research and Development Program of China (2023YFD1400400).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConflict of interest\u0026nbsp;\u003c/strong\u003eThe authors declare that they have no conflict of interest.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eOpen Access\u003c/strong\u003e This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the\u003c/p\u003e\n\u003cp\u003eoriginal author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article\u0026rsquo;s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article\u0026rsquo;s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.\u003c/p\u003e\u003cp\u003e\u003cstrong\u003ePublisher\u0026rsquo;s Note\u0026nbsp;\u003c/strong\u003eSpringer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eSupporting information\u0026nbsp;\u003c/strong\u003eAdditional supporting information may be found online in the Supporting Information section at the end of the article.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n \u003cli\u003eAcharya K, Tande C, Byamukama E (2016) Determination of \u003cem\u003eHeterodera glycines\u003c/em\u003e virulence phenotypes occurring in South Dakota. 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Hubei Agricultural Sciences 49:865-868. doi:10.14088/j.cnki.issn0439-8114.2010.04.044\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003eTable 1 Major QTLs associated with resistance to \u003cem\u003eHeterodera glycines\u003c/em\u003e (FI, CGR) and \u003cem\u003eM\u003c/em\u003e\u003cem\u003eeloidogyne\u0026nbsp;incognita\u003c/em\u003e (GI, RF, EMR, EGR) identified by Kruskal-Wallis and MQM analysis\u003c/p\u003e\n\u003cdiv align=\"\"\u003e\n \u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"100%\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eQTL name \u003csup\u003ea\u003c/sup\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eChr \u003csup\u003eb\u003c/sup\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 6px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eQTL region\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eMbp\u003csup\u003e\u0026nbsp;c\u003c/sup\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 6px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eSNP block\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"4\" valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eK\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e* Test1 (5,000 eggs in MI)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"4\" valign=\"top\" style=\"width: 20px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eK\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e* Test2 (10,000 eggs in MI)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"3\" valign=\"top\" style=\"width: 17px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eMQM Test1 (5,000 eggs in MI)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"3\" valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eMQM Test2 (10,000 eggs in MI)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eK\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e*\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eSignif. \u003csup\u003ed\u003c/sup\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eZYD \u003csup\u003ee\u003c/sup\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eSN\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eK\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e*\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eSignif.\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eZYD\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eSN\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eLOD \u003csup\u003ef\u003c/sup\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u003cstrong\u003ePVE (%)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eAdditive effect\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eLOD\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u003cstrong\u003ePVE\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e(%)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eAdditive effect\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eQTL-FI\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gs05-FI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e05\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e29.30-39.42\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e2726-2892\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e7.724\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e93.48\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e106.58\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.29\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e5.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e-7.85\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gm07a-FI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e07a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e0.00-7.60\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e3413-3531\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e6.691\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e123.85\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e104.27\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e5.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e9.50\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gs08a-FI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e08a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e9.96-14.77\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e4234-4471\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e12.37\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e******\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e83.31\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e105.90\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.14\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e4.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e-20.14\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gs09a-FI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e09a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e0.65-5.47\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e4833-4865\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e11.833\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e*****\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e85.78\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e106.37\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.44\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e5.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e-10.29\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gs20-FI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e20\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e0.48-2.51\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e10409-10447\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e5.731\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e87.26\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e106.34\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.14\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e4.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e-11.11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eQTL-CGR\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gs01b-CGR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e01b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e33.74-49.34\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e423-482\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e11.151\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e*****\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e53.05\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e65.50\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.51\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e5.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e-6.66\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gm02-CGR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e02\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e0.60-9.79\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e563-883\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e25.949\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e*******\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e83.13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e62.26\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e6.66\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e14.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e11.37\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gs03b-CGR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e03b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e36.91-45.51\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e1878-2080\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e11.847\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e*****\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e77.15\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e62.68\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e3.02\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e6.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e7.01\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gm11a-CGR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e11a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e0.02-10.12\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e5465-5804\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e14.522\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e******\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e75.34\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e62.42\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.99\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e6.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e6.30\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gm12-CGR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e12\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e3.91-12.82\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e6141-6256\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e13.226\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e******\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e76.70\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e62.89\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.73\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e6.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e7.03\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eQTL-GI\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.16\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e5.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi1-Gm01a-GI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e01a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e2.21-7.36\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e278-393\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e10.267\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e****\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e3.31\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e3.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.25\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e5.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e-0.12\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi1-Gs07a-GI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e07a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e3.90-24.25\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e3455-3925\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e9.756\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e****\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.89\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e3.17\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.62\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e6.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e-0.15\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi1-Gs13b-GI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e13b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e28.37-42.52\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e6863-7097\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e10.217\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e****\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.86\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e3.16\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e4.043\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e3.21\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e3.57\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.22\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e5.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e-0.15\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi1-Gs17-GI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e17\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e13.12-39.60\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e8857-9095\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e8.816\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e****\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.85\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e3.16\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e4.995\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e3.28\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e3.58\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.17\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e5.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e-0.22\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi1-Gs18b-GI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e18b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e48.54-57.75\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e9401-9567\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e4.661\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.95\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e3.15\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e11.291\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e*****\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e3.15\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e3.59\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.12\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e5.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.29\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi1-Gm19b-GI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e19b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e30.04-46.61\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e9696-10228\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e8.696\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e****\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e3.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e3.13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e13.577\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e******\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e4.02\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e3.52\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.16\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e5.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eQTL-RF\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi-Gm05b-RF\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e05b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e3.66-39.36\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e2523-2891\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e27.759\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e****\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e0.82\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e0.56\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e5.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi-Gm06c-RF\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e06c\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e49.54-51.13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e3366-3412\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e4.65\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e0.71\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e0.63\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.21\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e5.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.22\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi-Gs07a-RF\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e07a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e0.96-4.45\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e3432-3460\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e4.175\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e0.40\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e0.64\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e5.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.31\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi-Gm07c-RF\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e07c\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e9.62-17.90\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e3696-3904\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e9.522\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e****\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e1.09\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e0.62\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.26\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e5.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.24\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi-Gm10-RF\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e5.22-5.46\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e5149-5152\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e6.318\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e0.35\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e0.15\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.21\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e5.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi-Gm16a-RF\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e16a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e3.04-33.12\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e8380-8590\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e6.837\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e0.25\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e0.14\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e4.31\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e10.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.07\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eQTL-EMR\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi-Gm06b-EMR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e06b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e49.70-51.13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e3369-3412\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e6.139\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e23.49\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e18.61\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.05\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e5.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e4.45\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi-Gm16-EMR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e16\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e3.04-37.10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e8380-8683\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e10.88\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e*****\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e3.89\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.30\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e4.85\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e12.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e1.26\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi-Gm19a-EMR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e19a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e6.00-10.12\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e9639-9688\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e4.734\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e33.13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e18.56\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.75\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e10.44\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eQTL-EGR\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi-Gm05b-EGR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e05b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e3.66-39.36\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e2523-2891\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e14.273\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e******\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e4810.44\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e3640.31\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.35\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e6.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e542.74\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi1-Gm16b-EGR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e16b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e3.04-33.13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e8380-8590\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e9.006\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e****\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e589.61\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e307.03\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e5.75\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e14.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e177.03\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi-Gm19-EGR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e19\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e3.63-44.50\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e9635-10135\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e7.643\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e5469.18\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e3902.07\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 4px;\"\u003e\n \u003cp\u003e2.37\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 3px;\"\u003e\n \u003cp\u003e10.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7px;\"\u003e\n \u003cp\u003e2144.58\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 1px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n\u003c/div\u003e\n\u003cp\u003e\u003csup\u003ea QTL-FI or GI: QTLs related to female index (FI) or gall index (GI) phenotype. \u003cem\u003eHg4-Gs01-FI\u003c/em\u003e or \u003cem\u003eMi1-M01-RF.\u003c/em\u003e The name of QTL for FI or GI on Chr01 from \u003cem\u003eGlycine soja\u003c/em\u003e (Gs) ZYD00006 to \u003cem\u003eHeterodera glycines\u003c/em\u003e race 4 (HG Type 1.2.3.5.6.7) or \u003cem\u003eMeloidogyne incongnita\u003c/em\u003e race 1. Similarly, \u003cem\u003eHg4-Gm01-FI\u0026nbsp;\u003c/em\u003eor \u003cem\u003eMi1-Gm01-RF\u003c/em\u003e represents the name of QTL for FI or GI on Chr01 from \u003cem\u003eG. max\u003c/em\u003e cv. Suinong14 to \u003cem\u003eH. glycines\u003c/em\u003e race 4 or \u003cem\u003eM.incongnita\u003c/em\u003e race 1.\u0026nbsp;\u003c/sup\u003e\u003c/p\u003e\n\u003cp\u003e\u003csup\u003eb Chr: chromosome.\u0026nbsp;\u003c/sup\u003e\u003c/p\u003e\n\u003cp\u003e\u003csup\u003ec QTL region (Mbp) was from the first to the end physical position (SNP block).\u0026nbsp;\u003c/sup\u003e\u003c/p\u003e\n\u003cp\u003e\u003csup\u003ed Signif.: The significant P-values: \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05 (**), 0.01 (***), 0.005 (****), 0.001 (*****), 0.0005 (******). \u003cem\u003eK*\u003c/em\u003e score of \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05 (**) was shown in the table.\u003c/sup\u003e\u003c/p\u003e\n\u003cp\u003e\u003csup\u003ee The values of ZYD and SN show mean values of phenotype (FI or GI) related to ZYD00006 alleles or Suinong14 alleles.\u003c/sup\u003e\u003c/p\u003e\n\u003cp\u003e\u003csup\u003ef LOD scores and the percentage of phenotypic variance explained (PVE) are by means of MQM mapping analysis and the highest value of LOD and PVE values were shown in the SNP block in the QTL region. LOD (Logarithm of odds) \u0026gt; 2 was shown in the table. The additive effect with positive or negative values means increase or decrease in phenotypic values after the introgression segment of ZYD00006 alleles into Suinong14 background.\u003c/sup\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTable 2.\u003c/strong\u003e Common QTL intervals associated with resistance to \u003cem\u003eHeterodera glycines\u003c/em\u003e (HG) and \u003cem\u003eMeloidogyne incognita\u003c/em\u003e (MI) in a soybean chromosome segment substitution line (CSSL) population. Traits include female index (FI) and cysts per gram root (CGR) for SCN HG types 2.5.7 (race 5) and 1.2.3.5.6.7 (race 4), and galling index (GI), reproduction factor (RF), egg masses per gram root (EMR), and eggs per gram root (EGR) for \u003cem\u003eMI\u003c/em\u003e race 1\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\" width=\"100%\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 11px;\"\u003e\n \u003cp\u003eChr (LG) \u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 15px;\"\u003e\n \u003cp\u003eQTL interval\u003csup\u003e\u0026nbsp;b\u003c/sup\u003e\u003c/p\u003e\n \u003cp\u003eMbp\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 15px;\"\u003e\n \u003cp\u003eSNP block\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"3\" style=\"width: 58px;\"\u003e\n \u003cp\u003eQTL name\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 19px;\"\u003e\n \u003cp\u003eMI\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003eSCN4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 18px;\"\u003e\n \u003cp\u003eSCN5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e2 (D1b)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e8.80-9.79\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e854-883\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gm02-CGR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 18px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg5-Gs02a-CGR\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eHg5-Gs02-FI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e5 (A1)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e29.30-33.50\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e2726-2799\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi1-Gm05b-RF\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eMi1-Gm05b-EGR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gs05-FI\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 18px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e33.50-38.50\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e2799-2885\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi1-Gm05b-RF\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eMi1-Gm05b-EGR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gs05-FI\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 18px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg5-Gs05-FI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e38.50-39.36\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e2885-2891\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi1-Gm05b-RF\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eMi1-Gm05b-EGR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gs05-FI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 18px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg5-Gs05-FI\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eHg5-Gm05-CGR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e39.36-39.42\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e2891-2892\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gs05-FI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 18px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg5-Gs05-FI\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eHg5-Gm05-CGR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e7 (M)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e0.15-0.67\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e3414-3422\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gm07a-FI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 18px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg5-Gm07-CGR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e0.67-0.96\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e3422-3432\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gm07a-FI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 18px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg5-Gm07-CGR\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eHg5-Gs07b-FI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e0.96-2.50\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e3432-3441\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi1-Gs07a-RF\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gm07a-FI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 18px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg5-Gm07-CGR\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eHg5-Gs07b-FI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e2.50-4.45\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e3441-3460\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi1-Gs07a-RF\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gm07a-FI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 18px;\"\u003e\n \u003cp\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e10.81-15.50\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e3758-3788\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi1-Gm07c-RF\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 18px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg5-Gs07a-FI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e16 (J)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e32.40-33.13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e8552-8590\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi1-Gm16a-RF\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eMi1-Gm16b-EGR\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eMi1-Gm16-EMR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 18px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg5-Gs16-CGR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e33.13-36.50\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 15px;\"\u003e\n \u003cp\u003e8590-8666\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi1-Gm16-EMR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 18px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg5-Gs16-CGR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003csup\u003ea\u003c/sup\u003e Chr and LG was short for chromosome and linkage group.\u003c/p\u003e\n\u003cp\u003e\u003csup\u003eb\u003c/sup\u003e The major QTL region (\u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05** and LOD \u0026gt; 2) were shown in the table.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTable 3\u0026nbsp;\u003c/strong\u003eThe reported QTLs associated with resistance to \u003cem\u003eHeterodera glycines\u003c/em\u003e and \u003cem\u003eMeloidogyne incognita\u003c/em\u003e were mapped in chromosome segment substitution lines (CSSLs)\u003c/p\u003e\n\u003cdiv align=\"center\"\u003e\n \u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"100%\" class=\"fr-table-selection-hover\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 17px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eQTL name \u003csup\u003ea\u003c/sup\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 82px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eReported QTL (soybase.org)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd height=\"13\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd height=\"13\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 17px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eQTL-FI\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 82px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 0px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 17px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gs05-FI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 82px;\"\u003e\n \u003cp\u003eSCN 18-1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 0px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 17px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gm07a-FI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 82px;\"\u003e\n \u003cp\u003eSCN 46-1; SCN5-g16\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 0px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 17px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gs08a-FI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 82px;\"\u003e\n \u003cp\u003eSCN 8-4; SCN 9-1; SCN 9-3; SCN 29-5; SCN 30-3; SCN 39-8; SCN 42-1; SCN 45-4; SCN 50-1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 0px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 17px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gs09a-FI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 82px;\"\u003e\n \u003cp\u003eSCN5-g19\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 0px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 17px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gs20-FI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 82px;\"\u003e\n \u003cp\u003eSCN5-g52\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 0px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 17px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eQTL-CGR\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 82px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 0px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 17px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gs01b-CGR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 82px;\"\u003e\n \u003cp\u003eSCN 39-6; SCN 39-11; SCN5-g2; SCN4-g1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 0px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 17px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gm02-CGR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 82px;\"\u003e\n \u003cp\u003eSCN5-g4; SCN5-g3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 0px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 17px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gs03b-CGR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 82px;\"\u003e\n \u003cp\u003eSCN 33-7; SCN 44-15; SCN 51-1; SCN5-g6; SCN1-g1; SCN5-g5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 0px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 17px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gm11a-CGR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 82px;\"\u003e\n \u003cp\u003eSCN 46-6; SCN 49-1; SCN5-g22; SCN5-g25\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 0px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 17px;\"\u003e\n \u003cp\u003e\u003cem\u003eHg4-Gm12-CGR\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 82px;\"\u003e\n \u003cp\u003eSCN 10-4; SCN 10-5; SCN 11-1; SCN 11-2; SCN 39-4; SCN5-g27; SCN5-g26; SCN4-g9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 0px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 17px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eQTL-GI\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 82px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 0px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 17px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi1-Gs18b-GI\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 82px;\"\u003e\n \u003cp\u003eMi 1-3; Mi1-4; Mi 2-2; Mi 3-2; Mi 4-1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 0px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 17px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eQTL-RF\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 82px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 0px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 17px;\"\u003e\n \u003cp\u003e\u003cem\u003eMi-Gm10-RF\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 82px;\"\u003e\n \u003cp\u003eMi 1-1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 0px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n\u003c/div\u003e\n\u003cp\u003e\u003csup\u003ea QTL-FI or GI: QTLs related to female index (FI) or gall index (GI) phenotype. \u003cem\u003eHg4-Gs01-FI\u003c/em\u003e or \u003cem\u003eMi1-M01-RF.\u003c/em\u003e The name of QTL for FI or GI on Chr01 from \u003cem\u003eGlycine soja\u003c/em\u003e (Gs) ZYD00006 to \u003cem\u003eHeterodera glycines\u003c/em\u003e race 4 (HG Type 1.2.3.5.6.7) or \u003cem\u003eMeloidogyne incongnita\u003c/em\u003e race 1. Similarly, \u003cem\u003eHg4-Gm01-FI\u0026nbsp;\u003c/em\u003eor \u003cem\u003eMi1-Gm01-RF\u003c/em\u003e represents the name of QTL for FI or GI on Chr01 from \u003cem\u003eG. max\u003c/em\u003e cv. Suinong14 to \u003cem\u003eH. glycines\u003c/em\u003e race 4 or \u003cem\u003eM.incongnita\u003c/em\u003e race 1.\u0026nbsp;\u003c/sup\u003e\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":false,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Glycine max, G. soja, Soybean cyst nematode, Root-knot nematode, Transgressive inheritance, Chromosome segment substitution lines, QTL mapping","lastPublishedDoi":"10.21203/rs.3.rs-7587031/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7587031/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eSoybean cyst nematode (\u003cem\u003eHeterodera glycines\u003c/em\u003e, SCN) and southern root-knot nematode (\u003cem\u003eMeloidogyne incognita\u003c/em\u003e, RKN) are major limitation to soybean production, with few resistance sources available. Using 199 and 177 chromosome segment substitution lines (CSSLs) derived from \u003cem\u003eGlycine max\u003c/em\u003e cv. Suinong14 and \u003cem\u003eG. soja\u003c/em\u003e accession ZYD00006, we evaluated resistance to SCN HG type 1.2.3.5.6.7 (race 4) and RKN race 1, and compared the loci with those previously mapped for SCN HG type 2.5.7 (race 5). Despite both parents being susceptible, many CSSLs exhibited resistance levels exceeding both parents, while others showed extreme susceptibility, demonstrating strong transgressive segregation. QTL mapping identified 6 FI-associated and 9 CGR-associated loci for SCN race 4, and 17\u0026ndash;25 significant loci for RKN across two inoculum levels. Single resistance loci generally conferred only moderate effects, but the combination of multiple favorable alleles from both parents produced higher resistance. Comparative analysis revealed several shared QTL intervals among SCN races 4 and 5 and RKN race 1, suggesting partial common genetic control. These findings demonstrate that pyramiding multiple loci from wild and cultivated soybean can generate transgressive inheritance, producing resistant lines beyond parental performance and offering valuable targets for breeding durable, broad-spectrum nematode resistance.\u003c/p\u003e","manuscriptTitle":"Transgressive inheritance of broad-spectrum resistance to Heterodera glycines and Meloidogyne incognita in soybean chromosome segment substitution lines","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-10-09 13:07:55","doi":"10.21203/rs.3.rs-7587031/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"7b9ec5c0-79a7-4312-bb5c-e782f2b50e6b","owner":[],"postedDate":"October 9th, 2025","published":true,"recentEditorialEvents":[{"type":"decision","content":"Rejected","date":"2026-05-21T21:01:05+00:00","index":"","fulltext":""}],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2026-05-21T21:09:21+00:00","versionOfRecord":[],"versionCreatedAt":"2025-10-09 13:07:55","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-7587031","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7587031","identity":"rs-7587031","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}