Introduction
Materials and Methods
Plant cultivation and sample preparation
Characterization of phenotypic traits
Evaluation of glucosinolate contents
Free-radical scavenging activity
Statistical analysis
Results
Phenotypic traits of radish root
Glucosinolate contents of radish roots
Correlation analysis of phenotypic traits and glucosinolate profiles
Principal component analysis
Glucosinolate contents of root color-based groups
Discussion
Introduction
The edible roots of radish (Raphanus sativus L., 2n = 2x = 18), a representative of the Raphanus genus in the Brassicaceae family, are consumed worldwide. Radish sprouts and young leaves are also cooked, preserved by salting or pickling, or eaten in salads. According to Vavilov et al. (1926), radish originated from the eastern Mediterranean region and Middle East, and the current varieties might have been domesticated in India and other parts of Asia. Radish has been cultivated in Korea and China since 400 BC (Kaneko and Matsuzawa, 1993; Kurina et al., 2021; Vavilov et al., 1926). Historical reports of radish kimchi date back to the Three Kingdoms era (700 BC) in Korea, and traditional radish recipes have been published in at least 11 Korean recipe books within the past 100 years (Cho, 2010). Radish is one of the most widely cultivated vegetables in Korea and has been professionally bred for “four-season cultivation”.
In Asian countries, diverse varieties of radish are widely cultivated as large-rooted and long-season vegetables, whereas in the Americas and Europe, red, small-rooted, short-season radish varieties are cultivated. Black radish is used in salads in Spain because of its crispy texture, and in China, colorful radishes including 20-day red radish, green radish (green both outside and inside the root), and watermelon radish (green exterior and red interior) are consumed (Singh et al., 2017; Vavilov et al., 1926; Wang et al., 2020). In contrast to the colorful radishes cultivated in China, white radishes with green shoulders have been preferred in Korea for a long time because there is a perception that these types of radishes, especially egg-shaped ones, are of higher quality, being sweet and crunchy. Most radish and Brassica spp. researches conducted in Korea have focused on cultivation and disease resistance (Afroz et al., 2021; Geum et al., 2021; Kim et al., 2020; Kwon et al., 2020; Rajalingam et al., 2021), with few studies conducted on the color and appearance of radish roots (Singh et al., 2017). However, color is one of the most important traits reflecting quality, and affects consumer preference.
Glucosinolates (GSLs) are sulfur-containing secondary metabolites considered as the main health-promoting (anti-cancer, antioxidant, and anti-inflammatory) compounds in most cruciferous vegetables (Barillari et al., 2005). GSLs have been reported in various species and cultivars, as well as in specific plant organs (Hwang et al., 2019; Jeon et al., 2018; Rhee et al., 2020). Glucoraphasatin (GRH), which is derived from glucoerucin (GER), is abundant in radish (Fahey et al., 2001; Ishida et al., 2015; Kakizaki et al., 2017). Rhee et al. (2020) assessed the GSL distribution across the inner, middle, and outer leaves of kimchi cabbage (Brassica rapa L.), and Jeon et al. (2018) evaluated the GSL content at different vegetative growth stages in B. rapa (Jeon et al., 2018; Rhee et al., 2020).GSLs can be hydrolyzed by myrosinase to form breakdown products including isothiocyanates, which are responsible for the bitter taste of vegetables and exhibit cancer chemopreventive activity. Liu et al. (2018) compared nutritional values among different broccoli tissues and suggested various possible applications for broccoli by-products after analyzing sulforaphane derived from glucoraphanin (Liu et al., 2018). GRH is hydrolyzed by myrosinase when plant cells are disrupted to produce raphasatin; when derived from GRH, raphasatin was reported to induce more potent detoxification enzymes than other degradation products (Scholl et al., 2011). Raphasatin exhibits chemopreventive effects, including some toxic effects on human breast adenocarcinoma cells (Scholl et al., 2011; Suzuki et al., 2016; 2017), in which it induces apoptosis (Ibrahim et al., 2018). Radish also converts isothiocyanates from GSLs more efficiently than broccoli (De Nicola et al., 2013), with the addition of radish sprouts to broccoli sprouts promoting sulforaphane formation in the latter (Liang et al., 2018). Therefore, it is important to identify radish varieties that produce large amounts of GSLs, as they can provide health benefits to humans.
Montaut et al. (2010) provide an excellent summary of GSL analytical methods (Montaut et al., 2010). High-performance liquid chromatography (HPLC) has been used in many studies (Park et al., 2014) to analyze desulfated GSL with a UV-visible or photodiode array detector, but the desulfonation process is laborious, and some GSL may be insufficiently desulfonated at lower sulfatase concentrations. Recently, intact GSLs were identified and quantified using ultra- performance liquid chromatography (UPLC) with diode array detection-tandem mass spectrometry (MS/MS) in multiple reaction monitoring (MRM) mode (Gratacós-Cubarsí et al., 2010; Hwang et al., 2019). Several studies have analyzed GSL contents in the Brassicaceae family (Bhandari et al., 2015), but they often did not include GRH, which comprises 90% of all radish GSLs on average. Yi et al. (2016) investigated radish GSLs, but the numbers of germplasms and morphological traits analyzed were limited, and GSLs were analyzed in the form of desulfo-GSLs using HPLC (Yi et al., 2016).
Large-scale characterization and evaluation of the agronomic traits and health-promoting chemicals in radish germplasms would help identify new germplasm breeding materials. Therefore, in this study, we aimed to profile GSLs and characterize the phenotypes of radish germplasms. We evaluated the GSL contents of 110 radish germplasms using liquid chromatography (LC)-MS/MS in MRM mode, and characterized qualitative and quantitative traits. The data from the correlation analysis could promote the exploitation of favorable traits by breeders, as well as diet diversification given the health benefits associated with various radish germplasms.
Materials and Methods
Plant cultivation and sample preparation
Seeds of 110 radish germplasms were obtained from the National Agrobiodiversity Center (Jeonju, Korea), and seeds of the following 10 Korean commercial cultivars were purchased from various companies: Gwailmu (Con1; Asia Seed Co., Seoul, Korea), Meosjinmaskkalmu (Con2; Nongwoobio., Suwon, Korea), Taecheong (Con3; Syngenta, Iksan, Korea), Cheong- unmu (Con4; Farm Hannong, Seoul, Korea), Chorongmu (Con5; Farm Hannong), Mansa-hyeongtongmu (Con6; Nongwoobio.), Togwanggoldeumu (Con7; Farm Hannong), Baeksinaltari (Con8; Koregon, Seoul, Korea), Syupeogiljomu (Con9; Nongwoobio.), and Seohogoldeumu (Con10; Nongwoobio.; Appendix 1).
The seeds were sown in an experimental field containing compost and fertilizer at the end of August 2017. Plants were cultivated in the field following cultural practices recommended by the Rural Development Administration. Fertilizer (N-P-K-Ca-B = 60-40-60-75-1.5 ㎏/10 a) was applied before the seeds were sown. Radishes were harvested at 3-10 weeks after sowing (at the optimal growth stage) for phenotypic characterization, and all root samples were freeze-dried (LP500 vacuum freeze-drier; Ilshinbiobase Co., Seoul, Korea) directly at -70℃ for 1 week. The samples were then ground into fine powder and stored at -20℃ until further analysis.
Characterization of phenotypic traits
Phenotypic traits were characterized at full maturity in the field. Radish leaves and roots were examined for 5 qualitative and 10 quantitative traits based on modified descriptors from the International Union for the Protection of New Varieties of Plants (UPOV, 2021) and reference descriptors for Brassica and Raphanus (IBPGR, 1990).
The five qualitative traits were radish root peel color, root shoulder color, root flesh color, root shape, and the extent of root burial in soil. The 10 quantitative traits were total weight (Twe), root length (RL), root width (Rwi), root length-to- width ratio (RR), root weight (Rwe), leaf length (LL), leaf width (Lwi), leaf length-to-width ratio (LR), leaf weight (Lwe), and leaf number (LN). Each characteristic was examined using a digital caliper and digital balance. Three to five independent biological samples were examined to characterize the quantitative and qualitative phenotypic traits.
Evaluation of glucosinolate contents
GSLs were extracted following the method reported by Rhee et al. (2020). GSLs were isolated from 100-mg freeze-dried samples using 1 mL of solvent (Methanol: deionized water = 80:20, v/v). Then, each mixture was vortexed and centrifuged at 16,000 rpm and 4℃ for 10 min. The supernatant was transferred into a new tube, re-centrifuged, filtered using a 0.45-㎛ syringe filter, and diluted 10 times before an internal standard solution was added. Finally, the filtered solution was transferred into a brown vial for further analysis. For this experiment, the internal standard was prepared using 100 ppb glucotropaeolin. We analyzed 18 GSL standards (Phytoplan Diehm & Neuberger GmbH; Heidelberg, Germany) using a Waters Acquity UPLC system (Waters, Milford, MA, USA) coupled with a Xevo TQ-S system (Waters MS Technologies, Manchester, UK) and finally selected the following five GSLs to evaluate the GSL contents in radish germplasms (Appendix 4): Glucoraphasatin (GRH), glucoraphenin (GRE), glucobrassicin (GBR), Glucoerucin (GER), and glucoberteroin (GBE). Glucotropaeolin was used as the internal standard to identify and quantify the GSLs in 110 germplasms and 10 commercial cultivars. Chromatographic separation was carried out using an Acquity UPLC BEH C18 (1.7 ㎛, 2.1 × 100 ㎜) column (Waters Corp., Manchester, UK). The flow rate was kept at 0.25 mL/min, the column temperature was maintained at 35℃, and the injection volume was 5 μL. The mobile phase comprised 0.1% trifluoroacetic acid in distilled water (A) and 0.1% trifluoroacetic acid in methanol (B). The UPLC gradient conditions were as follows: initial condition, 100% A; 0.0-1.0 min, 100% to 95% A; 1.0-4.0 min, 95% to 0% A; 4.0-4.5 min, 0% to 100% A; 4.5-5 min, 100% A; 5-15 min, 100% A. The mass spectrometry instrument was operated in negative ion electrospray ionization (ESI) and MRM modes. Data were acquired using MassLynx 4.1 software. GSLs were identified by comparing their retention times and MS and MS/MS fragmentation spectra with those of commercial standards. Each MRM transition was set as follows {compound name (retention time, parent molecular weight > daughter transition weight)}: GRH (5.18 min, 417 > 175.69), GRE (6.21 min, 433 > 418.5), GER (4.87 min, 419 > 177.71), GBE (6.21 min, 433 > 127.78), GBR (5.51 min, 446 > 204.69), and glucotropaeolin (4.75 min, 407 > 165.94). The final concentration of each GSL was calculated based on its curve area relative to that of the internal standard (glucotropaeolin) and linear regression equations derived from the calibration curve of the corresponding standard. The final concentrations of individual GSLs are presented in units of ㎍/g sample dry weight (DW).
Free-radical scavenging activity
To evaluate antioxidant activity in the radish germplasms, we modified the 2,2-diphenyl-1-picrylhydrazyl (DPPH) test protocol of Brand-Williams et al. (1995). DPPH powder (Sigma-Aldrich, St. Louis, MO, USA) was dissolved in 200 mL methanol to make a 150 mM DPPH solution, which was shaken in the dark for 1 h. Then, 1 mL of 80% methanol was added to each 2-mg freeze-dried sample, and the mixture was sonicated for 1 h and then centrifuged. The supernatant was analyzed for antioxidant activity. For the analysis, 150 μL of 150 mM DPPH and 100 μL of 2,000 ppm sample extract were mixed. DPPH solution and ascorbic acid (Sigma-Aldrich, Milwaukee, WI, USA) were used as a blank and standard, respectively. The absorbance at a wavelength of 517 ㎚ was measured. Experiments were performed in triplicate with independent germplasms. The free-radical scavenging activity was calculated as follows:
DPPH free-radical scavenging activity (%) = [1 - (A sample - A sample blank) / (A control - A control blank)] × 100%, where A is the absorbance at 517 ㎚.
Statistical analysis
All experiments of GSL contents were conducted in technical triplicate, and the average ± standard deviation of GSL contents were calculated (㎍/g of root DW). Correlation analysis of the 10 phenotypic traits and individual and total compound amounts, as well as principal component analysis (PCA) of GSL contents and free-radical scavenging activities, were performed using XLSTAT software (Addinsoft, Paris, France). Analysis of variance followed by Duncan’s multiple range test {least significant range (LSR), p < 0.05} was performed to determine if the content of each compound varied significantly by phenotypic traits. Student’s t-test was done to identify significant-phenotypic differences between germplasms and cultivars.
Results
Phenotypic traits of radish root
In total, 15 phenotypic traits important for radish breeding were evaluated, including root color, shape, weight, and length. Five qualitative and ten quantitative traits of 110 radish germplasms and 10 Korean commercial cultivars were assessed (Appendix1 and 2), and the results are summarized in Table 1 and 2. Broad variation in all qualitative traits was observed in the radish germplasms compared to the commercial cultivars. In the latter, there was relatively little variation, especially in root peel color and root shoulder color. The root peel color of most germplasms (70.9%) and all cultivars (100%) was white. For the rest of the germplasms, two other root peel colors, bronze-green and red, were dominant. The radish germplasms produced roots with various root shoulder colors, including green (31.8%), whereas only the green shoulder color was observed in the cultivars. The root flesh color in the germplasm-produced roots and cultivars was mainly white (77.3% and 90%, respectively). Nineteen germplasms, but no cultivars, produced roots with green flesh. The germplasms produced various root shapes, such as a narrow rectangle (31.8%), sphere (19.1%), and rectangle (17.3%), whereas the roots of the cultivars were mostly elliptical (70%) or gourd- shaped (20%). Regarding the root position in soil, germplasm- produced roots were mostly buried (46.4%) or half buried (30.9%), whereas most cultivar-produced roots were half buried in soil (60%).
Table 1.
Quantitative traits were more variable in the radish germplasms compared to the cultivars, especially Twe, Rwe, Lwe, RL, LL, Rwi, Lwi, RR, LR, and LN (Table 2). The average Twe of the cultivars was higher (2,088.9 g) than that of the germplasms (1,827.5 g), as was the average Rwe (1,753.3 vs. 1,286.3 g). Conversely, the average Lwe of the germplasms was higher (541.1 g) than that of the cultivars (335.3 g), and the average RL and LL were greater in the germplasms (24.1 and 45.6 ㎝, respectively) than in the cultivars (20.6 and 39.9 ㎝, respectively), despite the average Rwi (11.8 vs. 10 ㎝) and Rwe being greater in the latter. Both Rwe and Lwe were significantly different between germplasms and cultivars (p < 0.05) in Table 2. Leaves derived from the germplasms were wider (average width, 16.7 ㎝) than in the cultivars (average, 15.9 ㎝), but the LR was similar between the germplasms (2.8) and cultivars (2.5). For the RR, there was a marked difference between the germplasms (2.7) and cultivars (1.7). The numerical difference in the ratio was small, but the RR range of the germplasms was much broader than that of the cultivars. The average LN was similar between the germplasms (26.5) and cultivars (25.1), with the former having a broader range (8.7-59) compared to the latter (19-33).
Table 2.
No. | Phenotypic traits | Germplasms | Cultivars | |||||
Range | Median | Average ± SDz | Range | Median | Average ± SDz | |||
1 | Total weight (Twe; g) | 45.4 – 3420.0 | 1799.1 | 1827.5 ± 637.8 | 723.3 - 2776.7 | 2113.3 | 2088.7 ± 621.2 | |
2 | Root weight (Rwe; g)*y | 45.0 - 2918.3 | 1280.0 | 1286.3 ± 463.8 | 588.3 - 2363.3 | 1948.3 | 1753.3 ± 529.4 | |
3 | Leaf weight (Lwe; g)*y | 0.4 - 1833.3 | 515.2 | 541.1 ± 279.0 | 135.0 - 575 | 363.3 | 335.3 ± 129.5 | |
4 | Root length (RL; ㎝) | 2.5 – 48.0 | 23.2 | 24.1 ± 10.8 | 12.0 - 25.8 | 21.2 | 20.6 ± 4.6 | |
5 | Root width (RW; ㎝) | 2.1 - 41.8 | 9.6 | 10.0 ± 3.8 | 10.5 - 12.6 | 11.8 | 11.8 ± 0.6 | |
6 | Leaf length (LL; ㎝) | 14.2 – 62.0 | 46.8 | 45.6 ± 8.9 | 32.3 - 48.3 | 38.5 | 39.9 ± 4.6 | |
7 | Leaf width (LW; ㎝) | 8.4 - 25.8 | 17.1 | 16.7 ± 3.3 | 11.8 - 18.9 | 17.3 | 15.9 ± 2.3 | |
8 | Root ratio (RR; len/wid) | 0.7 - 6.2 | 2.3 | 2.7 ± 1.5 | 1.1 - 2.1 | 1.9 | 1.7 ± 0.3 | |
9 | Leaf ratio (LR; len/wid) | 1.7 - 4.1 | 2.8 | 2.8 ± 0.5 | 2.0 - 3.1 | 2.5 | 2.5 ± 0.4 | |
10 | Leaf number (LN) | 8.7 – 59.0 | 25.3 | 26.5 ± 11.0 | 19.0 – 33.0 | 25.4 | 25.1 ± 4.4 |
Glucosinolate contents of radish roots
The total GSL content and content of each of the five individual GSLs examined in the radish germplasms and cultivars (Appendix 3) are summarized in Table 3. The average total GSL content in the germplasm-produced roots was 7,535.7 ㎍/g DW. The average content of GRH, the main constituent (73.2%) of the total GSL content in radish, was 5,512.9 ㎍/g DW. The GRH content also varied the most widely among the five GSLs (123.8-12,922.0 ㎍/g DW). The average contents of GRE, GER, GBR, and GBE in the germplasm- produced roots were 1,716.9, 165.8, 127.1, and 12.9 ㎍/g DW, respectively. The GBE, GBR and GER contents did not differ significantly from each other, whereas the GRE, GRH, and total GSL contents differed significantly from the contents of the other constituents (Duncan’s LSR, p < 0.05). The GBE, GBR, GER, and GRE contents were similar to each other in the cultivar-produced roots, whereas the GRH and total GSL contents were significantly different from the contents of the other constituents (Duncan’s LSR, p < 0.05). The GRE and GBE contents were significantly different between germplasm-and cultivar-produced roots (Student’s t-test, p < 0.05) in Table 3.
Table 3.
110 germplasms | 10 commercial varieties | ||||||
Average | Range | Top germplasm | Average | Range | Top variety | ||
Total glucosinolate content | 7535.7dy | 154.3 - 18438.5 | 299453 | 7989.6cy | 4093.5 - 10873.1 | Seohogoldeumu | |
Glucoraphasatin | 5512.9cy | 123.8 - 12922.0 | 215011 | 6794by | 2531.3 - 9247 | Seohogoldeumu | |
Glucoraphenin*z | 1716.9by | 26.3 - 5653.2 | 306869 | 956.8ay | 509.6 - 1307.1 | Syupeogiljomu | |
Glucoerucin | 165.8ay | 2.4 - 534.5 | 299453 | 155.4ay | 40.9 - 292.9 | Gwailmu | |
Glucobrassicin | 127.1ay | 0.7 - 1243.9 | 306869 | 73.6ay | 7.9 - 268.4 | Seohogoldeumu | |
Glucoberteroin*z | 12.9ay | 0 (ND) - 59.1 | 299453 | 9.8ay | 4.7 - 17.3 | Seohogoldeumu |
zAsterisk indicates significant differences in the content of a given compound between germplasms and cultivars (t-test, p < 0.05).
yDifferent characters indicate significant differences in mean content (Duncan’s multiple comparison test, p < 0.05) for each germplasm or cultivar. The glucoraphenin and glucoberteroin contents differed significantly between germplasms and cultivars. No significant differences were found for the other GSLs.
The roots of the following germplasms contained the highest amounts of the specified GSLs (Table 3): IT215011, GRH; IT306869, GRE and GBR; IT299453, GER and GBE. Among the 10 cultivars, Seohogoldeumu had the highest total GSL content (and the highest GRH, GBR, and GBE contents), while Syupeogiljomu had the highest GRE content and Gwailmu had the highest GER content. The content range for all GSLs was broader in germplasm-produced roots than Korean cultivar-produced roots.
Correlation analysis of phenotypic traits and glucosinolate profiles
Correlation analysis of the 15 phenotypic traits and contents of the individual GSLs was performed using XLSTAT software. The Pearson correlation coefficients are presented in Table 4. Twe, which was calculated by summing Rwe and Lwe, was more strongly correlated with Rwe (r = 0.916) than with Lwe (r = 0.725). Twe was also significantly correlated with LL (r = 0.651), RL (r = 0.580), and LN (r = 0.529). Rwe was significantly correlated with RL (r = 0.585), and Lwe was significantly correlated with LL (r = 0.684) and LN (r = 0.629). The RR was more strongly correlated with RL (r = 0.918) than with Rwi (r = -0.463). The RL was negatively correlated with DPPH antioxidant scavenging capacity (r = -0.431). LL was significantly correlated with Lwi (r = 0.678).
Table 4.
Traits | Twez | Rwe | Lwe | RL | Rwi | RR | LL | Lwi | LR | LN | GRH | GRE | GBR | GER | GBE | TG |
Rwey | 0.92**g | |||||||||||||||
Lwex | 0.73**g | 0.39**g | ||||||||||||||
RLw | 0.58**g | 0.59**g | 0.33**g | |||||||||||||
Rwiv | 0.32**g | 0.37**g | 0.11 | -0.16 | ||||||||||||
RRu | 0.33**g | 0.28**g | 0.28**g | 0.92**g | -0.46**g | |||||||||||
LLt | 0.65**g | 0.47**g | 0.68**g | 0.14 | 0.30**g | 0.01 | ||||||||||
Lwis | 0.48**g | 0.35**g | 0.49**g | 0.07 | 0.23*gh | -0.02 | 0.68**g | |||||||||
LRr | 0.25**g | 0.18 | 0.26**g | 0.10 | 0.11 | 0.03 | 0.45**g | -0.33** | ||||||||
LNq | 0.53**g | 0.34**g | 0.63**g | 0.45**g | -0.05 | 0.42**g | 0.25**g | -0.06 | 0.38**g | |||||||
GRHp | 0.35**g | 0.30**g | 0.29**g | 0.21*gh | 0.07 | 0.16 | 0.27**g | 0.18 | 0.13 | 0.22*gh | ||||||
GREo | 0.18 | 0.23*gh | 0.02 | 0.06 | 0.17 | -0.02 | 0.17 | 0.16 | 0.04 | -0.13 | 0.76**g | |||||
GBRm | -0.11 | -0.09 | -0.10 | -0.20*gh | 0.06 | -0.17 | 0.07 | 0.08 | 0.02 | -0.13 | 0.49***f | 0.59**g | ||||
GERl | 0.32**g | 0.31**g | 0.20*gh | 0.23*gh | 0.06 | 0.17 | 0.19*gh | 0.10 | 0.12 | 0.20*gh | 0.94**g | 0.74**g | 0.49**g | |||
GBEk | 0.18 | 0.17 | 0.11 | 0.12 | 0.03 | 0.11 | 0.20*gh | 0.14 | 0.10 | 0.01 | 0.86**g | 0.76**g | 0.55**g | 0.89**g | ||
TGj | 0.31**g | 0.29**g | 0.23*gh | 0.17 | 0.10 | 0.12 | 0.25**g | 0.18 | 0.11 | 0.13 | 0.98**g | 0.86**g | 0.57**g | 0.93**g | 0.88**g | |
DPPHi | -0.39**g | -0.35**g | -0.29**g | -0.43**g | -0.03 | -0.38**g | -0.26**g | -0.17 | -0.16 | -0.28**g | -0.10 | -0.01 | 0.17 | -0.15 | -0.10 | -0.08 |
Bold characters: r > 0.6, p < 0.01. zTwe: total weight (g), yRwe: root weight (g), xLwe: leaf weight (g), wRL: root length (㎝), vRwi: root width (㎝), uRR: root length-to-width ratio, tLL: leaf length (㎝), sLwi: leaf width (㎝), rLR: leaf length-to-width ratio, qLN: leaf number, pGRH: glucoraphasatin, oGRE: glucoraphenin, mGBR: glucobrassicin, lGER: glucoerucin, kGBE: glucoberteroin, jTG: total GSL content, including GRH, GRE, GBR, GER, and GBE, iDPPH: 2,2-diphenyl-1-picrylhydrazyl. h*, g** and f*** indicate significant correlations between two traits at p < 0.05, p < 0.01 and p < 0.001, respectively.
In Table 4, GRH was significantly correlated with the other GSLs, except GBR. GRH, the most abundant GSL in radish root, was highly correlated with the total GSL content (r = 0.984) and GER (r = 0.938). The strongest correlation among the five GSLs was observed between GRH and GER (r = 0.938), in accordance with previous reports. GRH was also strongly correlated with GBE (r = 0.858) and GRE (r = 0.755). GER (r = 0.934) and GBE (r = 0.878) were more strongly correlated with the total GSL content than was GRE (r = 0.755). GBR had weaker correlations with the other GSLs but was the only GSL positively correlated with DPPH anti-oxidant scavenging (r = 0.166).
Principal component analysis
Principal component analysis was conducted to analyze the relationships among GSL profiles (total GSL content, GRH, GBR, GER, GBE, and GRE) and antioxidant activity (DPPH scavenging) in the 110 radish germplasms (Fig. 1). The first two principal components (F1 and F2), represented as the x- and y-axis in the PCA biplot (Fig. 1), explained 68.88% and 15.97% (sum, 84.85%) of the total variance, respectively.
Five components, namely the total GSL content (20.1%), GRH (18.8%), GER (18.5%), GBE (17.8%), and GRE (15.9%), were the main contributors to F1, whereas DPPH activity (78.8%) and GBR (17.5%) were the main contributors to F2. The six GSL components had positive loadings on F1, with the total GSL content and GRH having the highest loadings, whereas DPPH activity was negatively loaded. DPPH activity, GBR, and GRE had positive loadings on F2, and the total GSL content, GBE, GRH, and GER had negative loadings (Fig. 1).
Based on their location in the PCA plot, the GSL contents may share some common tendencies. Along F1, more positive values reflect higher total GSL, GRH, GER, GBE, and GRE contents, and vice versa. These components do not make a large contribution to F2. More positive values for both F1 and F2 reflect higher GBR content. When the F1 value is positive and the F2 value is negative, the GBR content is low. When the F1 value is negative, the GBR content is intermediate. A negative F1 value and positive F2 value reflects high DPPH activity. In contrast to F2, DPPH activity does make a large contribution to F1, so observations located in the upper half of the plot indicate high DPPH activities and vice versa.
In the PCA biplot, the 110 germplasms were separated into five groups (Fig. 1). Group I (circled in red) contains five radish germplasms (4.5%) and is located in the upper right quadrant between +3.9 and +6.5 on the F1 axis and -0.4 and +2.8 on the F2 axis. The germplasms in Group I had > 10,000 ㎍/g DW GRH, > 15,000 ㎍/g DW for the total GSL content, > 100 ㎍/g DW GBR, and relatively high total GSL and GRH contents compared to the other groups. Two accessions, IT299453 (No. 104) and IT306869 (No. 107), had high total GSL content (18,438 and 18,018 ㎍/g, respectively) but exhibited low and moderate levels of DPPH activity, respectively. IT299453 (No. 104) was also located below IT306869 (No. 107). The IT215011 (No. 36) germplasm had the highest GRH content (12,922 ㎍/g DW) but only a moderate GBR content, and also exhibited moderate DPPH activity. It is positioned on the left side (negative F2) of the plot. Both IT306869 (No. 107) and IT215011 (No. 36) had high total GSL and GRH contents, with moderate levels of DPPH activity.
Group II contains three radish germplasms (2.7%) and is located in the upper right quadrant of the plot, within +0.9 to +1.7 on the F1 axis and +1.5 to +2.8 on the F2 axis (Fig. 1). The germplasms in Group II had total GSL contents of > 9,500 ㎍/g DW and antioxidant activities of > 40%. The GRE and GBR contents were moderate to high. The GSL contents were lower compared to Group I, but the antioxidant activities were higher. Of the three germplasms, IT262036 (No. 71) had the highest GBR content (533.3 ㎍/g DW), and its position indicates an association with GBR-related variables. IT297120 (No. 100) also had a high GBR content (449.1 ㎍/g DW), but its total GSL content was low (although its DPPH activity was still high). IT250790 (No. 53) had a high GRE content (4,421.2 ㎍/g DW).
Group III contains three germplasms and is located within -1.3 to -0.2 on the F1 axis and +2.3 to +3.9 on the F2 axis (Fig. 1). The germplasms in Group III exhibited high antioxidant activities (> 40%, with the highest activity level being 63.1%), moderate-to-low GSL contents, and high GBR contents (> 100 ㎍/g DW). Of the three germplasms, IT264178 (No. 80) exhibited the highest DPPH antioxidant activity but had a low GSL content; IT278682 (No. 84) had the highest GBR content (591.2 ㎍/g) in the group.
Group IV contains 34 germplasms, and is located within -3 to -1.7 on the F1 axis and -1.8 to +3.0 on the F2 axis (Fig. 1). Group IV members had low individual and total GSL contents. Group IV was further divided into three subgroups (Groups i- iii) with 3, 28, and 3 accessions, respectively, according to the antioxidant activity level. Group i members exhibited the highest antioxidant activities (> 45%), followed by Group ii (15-38%) and Group iii members (< 13%). The antioxidant activity levels in Group i were high, although the GSL contents were not. IT278727 (No. 88), IT136498 (No. 22), and IT208400 (No. 31) produced radish roots with red peel. IT208400 (No. 31) exhibited the highest antioxidant activity (59.8%) despite having a low total GSL content (1,334 ㎍/g DW). Group ii members had low GSL contents and exhibited largely similar levels of antioxidant activity. Group iii members {IT103811 (No. 14), IT112253 (No. 16), and IT278685 (No. 87)} exhibited the lowest antioxidant activities (< 13%) and did not produce red roots. IT278685 (No. 87) exhibited the lowest antioxidant activity (4.3%) and had the lowest GSL content.
Group V is positioned in the center of the plot, within -1.7 to +3.5 on both the F1 and F2 axis, and contains 65 germplasms (Fig. 1). No individual traits obviously characterized the group members, and the values of most traits were in the moderate range. To distinguish this group more precisely, more traits need to be analyzed.
Glucosinolate contents of root color-based groups
We classified radish germplasms based on their root color phenotypes, as follows: color group 1, red peel with a red shoulder and white flesh; color group 2, white peel with a green shoulder and red flesh; color group 3, bronze-green peel and shoulder with green flesh; color group 4, white peel, shoulder, and flesh (totally white); and color group 5, white peel with a green shoulder and white flesh (Fig. 2).
Color group 1 included 13 germplasms with a mean total GSL content of 4,424.2 ㎍/g DW (range: 642.0-13,044.0 ㎍/g DW; Fig. 2), which was the lowest mean content among all color groups. Color group 2 included six germplasms with a mean total GSL content of 7,082.3 ㎍/g DW (range: 2,660.9- 11,859.5 ㎍/g DW). Color group 3 included 19 germplasms with a mean total GSL content of 9,864.9 ㎍/g DW (range: 1,532.8-18,438.5 ㎍/g DW), which was the highest mean content among all color groups. Color group 4 included 43 germplasms with a mean total GSL content of 7,356.2 ㎍/g DW (range: 154.3-18,017.9 ㎍/g DW), and color group 5 included 29 germplasms with a mean total GSL content of 7,764.3 ㎍/g DW (range: 274.4-16,073.8 ㎍/g DW).
Images above the bar graph show the exterior and interior of the roots in each group with the highest GSL contents (IT102560, IT262036, IT299453, IT306869, and IT215011). In color group 1, the mean GRH, GRE, GBR, GER, and GBE contents were, respectively, 3,214.3, 10,613, 62.43, 81.13, and 5.23 ㎍/g DW, comprising 72.7%, 24%, 1.4%, 1.8%, and 0.1% of the total GSL content. Of the five color groups, group 1 germplasms had the lowest individual GSL contents, but the antioxidant activity was not as low. We further divided this group into two subgroups based on a root weight cutoff of 250 g. Of 13 germplasms, four weighed less than 250 g; these small radishes are eaten in salads or as pickles in Europe and the United States. The small radishes had lower GSL contents compared to the bigger radishes, but the antioxidant activities were high.
In color group 2, the mean GRH, GRE, GBR, GER, and GBE contents were, respectively, 4,928.7, 1,690.7, 364.9, 90.8, and 7.2 ㎍/g DW, comprising 69.6%, 23.9%, 5.2%, 1.3%, and 0.1% of the total GSL content (Fig. 2). The GBR content was significantly higher than in the other groups, and the mean antioxidant activity was the highest. In color group 3, the mean GRH, GRE, GBR, GER, and GBE contents were, respectively, 6,807.1, 2,560.2, 253.9, 219.7, and 24 ㎍/g DW, comprising 69%, 26%, 2.6%, 2.2%, and 0.2% of the total GSL content. The individual GSL contents were significantly higher than in the other groups, but unexpectedly, the mean antioxidant activity (DPPH free-radical scavenging activity) was not high.
In color group 4, the mean GRH, GRE, GBR, GER, and GBE contents were, respectively, 5,361.1, 1,732.7, 95.2, 172, and 11.4 ㎍/g DW, comprising 72.9%, 23.6%, 1.1%, 2.3%, and 0.2% of the total GSL content. In color group 5, the mean GRH, GRE, GBR, GER, and GBE contents were, respectively, 6,041.3, 1,440.4, 95.2, 174.8, and 12.6 ㎍/g DW, comprising 77.8%, 18.6%, 1.2%, 2.3%, and 0.2% of the total GSL content (7,764.3 ㎍/g DW). Radishes belong to this group are known as kimchi radishes and are particularly preferred in Korea.
Discussion
We found that various phenotypic traits were highly correlated with each other. In cases where desirable and undesirable traits are highly correlated, breeding programs employ selection processes to uncouple the correlations. According to Jatoi et al. (2011), Twe was highly correlated with Lwi (r = 0.70) and RL (r = 0.71) in their radish samples. In this study, we observed a significant correlation between Twe and LL (r = 0.65), as did Jatoi et al. (2011). Color is an important external characteristic used to evaluate radish quality. Customers in Korea prefer white radishes with green shoulders. This type of radish has a sweet taste and crispy texture, and is popularly used to make kimchi, a traditional Korean side dish, along with soups and various other side dishes. Consumers often modify their dietary habits to improve health. This could explain the recent increase in popularity of food products derived from Brassica vegetables, and the development of experimental products such as broccoli puree with lactic acid (Cai et al., 2019) and muffins enriched with dietary fiber from kimchi (Heo et al., 2019). Brassica plants contain GSLs that can be degraded by intestinal microorganisms to produce bioactive metabolites such as isothiocyanates, which have anti-cancer and other biological properties (Aires et al., 2009). In this study, five GSLs, four aliphatic GSLs (GER, GRH, GRE, and GBE) and one indole GSL (GBR) were screened and quantified in 110 radish germplasms. Three aliphatic GSLs (GRH, GER, and GBE) were significantly correlated with one another but not with GBE or the indole GSL, GBR. This could be due to differences in biosynthetic pathways and precursors, with GRH, GER, and GBE sharing a similar 4C pathway, whereas GBE is derived from a 5C pathway. GBR, the indole GSL, has a different amino-acid precursor, tryptophan.
The GSL contents in radish were found to be correlated with some phenotypic traits. The total GSL content was significantly correlated with LL but not with RL. A similar observation was reported for kimchi cabbage by Jeon et al. (2018) and Kakizaki et al. (2017), who studied the synthesis and movement of GSL (Jeon et al., 2018; Kakizaki et al., 2017). Yi et al. (2016) reported no strong correlations among root shape, pithiness, sweetness, peel color, length, and GSL content (Yi et al., 2016). Root phenotypic characters such as color, shape, and length were not strongly associated with the GSL profile of the radish germplasms in this study, implying that the root phenotype does not reflect GSL contents. However, based on this finding, it may be possible to develop radish varieties of various phenotypes with high GSL contents.
According to our PCA, GSL contents were not strongly correlated with DPPH activity. Raphasatin, a degradation product of GRH, was reported to more potently induce detoxification enzymes than other degradation products (Scholl et al., 2011; Suzuki et al., 2016). A more complete understanding of the overall antioxidant effect could be achieved by measuring the antioxidant activities of isothiocyanates such as raphasatin. The germplasms in Group III in the PCA plot in Fig. 1 produce small red radishes with high DPPH free-radical scavenging activities. To determine whether the antioxidant effect is influenced by the root size or red pigment, small non-red radish germplasms can be analyzed. Red radishes contain higher levels of high anthocyanins and other phenolic compounds than non-pigmented radishes (Singh et al., 2017). Hence, they can be used to prepare healthful, nutrient-dense dishes and nutraceutical formulations.
In this study, we assessed various phenotypic traits, DPPH free-radical scavenging activity, and the contents of five GSLs in radish germplasms. The results regarding GSL levels in radishes, and their relationships with leaf and root characteristics, could be used as baseline data by breeders and nutraceutical companies. Moreover, the biochemical and phenotypic information provided by this study may encourage consumers to diversify their eating habits.
Appendix
Appendix 1.
No. | IT No. | Root peel color | Root shoulder color | Root fresh color | Root shape | Root position in soil |
1 | 32729 | White | Green | White | N-rectangle | Half B |
2 | 100590 | White | Green | White | Rectangle | Half B |
3 | 100649 | White | Green | White | N-rectangle | Half B |
4 | 100678 | White | Green | White | Rectangle | Mostly B |
5 | 100684 | White | Green | White | Almond | Mostly B |
6 | 100689 | White | Green | White | Egg | Half B |
7 | 100691 | White | Green | White | Rectangle | Mostly B |
8 | 100695 | White | Green | White | Rectangle | Half B |
9 | 102376 | White | Green | White | N-rectangle | Mostly B |
10 | 102378 | White | Green | White | N-rectangle | Half B |
11 | 102395 | White | Green | White | N-rectangle | Half B |
12 | 102560 | Red | Red | White | Egg | Half B |
13 | 103802 | White | Green | White | Rectangle | Mostly B |
14 | 103811 | White | Green | White | Oval | Mostly B |
15 | 104055 | White | Green | White | W-rectangle | Mostly B |
16 | 112253 | White | White | White | N-rectangle | Mostly B |
17 | 112255 | White | White | White | N-rectangle | Mostly B |
18 | 112257 | White | White | White | N-rectangle | Mostly B |
19 | 112258 | White | White | White | N-rectangle | Mostly B |
20 | 119000 | White | Green | White | Rectangle | Mostly B |
21 | 136485 | White | White | White | N-rectangle | Mostly B |
22 | 136498 | Red | Red | White | Spherical | Mostly A |
23 | 166993 | White | White | White | Rectangle | Mostly B |
24 | 166995 | White | White | White | N-rectangle | Half B |
25 | 166997 | White | White | White | N-rectangle | Mostly B |
26 | 185738 | White | Green | White | Egg | Mostly B |
27 | 188102 | Bronze-green | Bronze-green | Green | W-rectangle | Mostly A |
28 | 203316 | Bronze-green | Bronze-green | Green | W-Oval | Mostly A |
29 | 203531 | White | Green | White | Rectangle | Mostly B |
30 | 204160 | White | Green | White | Rectangle | Mostly B |
31 | 208400 | Red | Red | White | Egg | Mostly A |
32 | 209937 | Bronze-green | Bronze-green | Green | Egg | Mostly A |
33 | 209974 | White | White | White | N-rectangle | Half B |
34 | 210203 | White | White | White | N-rectangle | Mostly B |
35 | 213153 | White | Green | White | Egg | Mostly B |
36 | 215011 | White | Green | White | Gourd | Mostly B |
37 | 215079 | White | White | White | N-rectangle | Half B |
38 | 218925 | White | Green | White | Egg | Mostly B |
39 | 220675 | Bronze-green | Bronze-green | Green | Egg | Mostly A |
40 | 221952 | Bronze-green | Bronze-green | Green | Rectangle | Mostly A |
41 | 221955 | White | White | White | Almond | Mostly B |
42 | 221956 | White | White | White | Almond | Mostly B |
43 | 221958 | White | White | White | Almond | Half B |
44 | 221959 | White | White | White | Rectangle | Half B |
45 | 223576 | Bronze-green | Bronze-green | Green | W-rectangle | Above |
46 | 228857 | Bronze-green | Bronze-green | Green | Egg | Above |
47 | 228870 | Bronze-green | Bronze-green | Green | Spherical | Above |
48 | 250738 | White | White | White | N-rectangle | Mostly B |
49 | 250765 | White | White | White | N-rectangle | Mostly B |
50 | 250775 | Bronze-green | Bronze-green | Green | Spherical | Above |
51 | 250777 | Bronze-green | Bronze-green | Green | Egg | Above |
52 | 250788 | White | White | White | N-rectangle | Mostly B |
53 | 250790 | White | White | White | W-rectangle | Mostly B |
54 | 250792 | Bronze-green | Bronze-green | Green | Rectangle | Mostly A |
55 | 250794 | Bronze-green | Bronze-green | Green | Rectangle | Mostly A |
56 | 261944 | Red | Red | White | Spherical | Half B |
57 | 261947 | White | White | White | N-rectangle | Half B |
58 | 261953 | White | White | White | Spherical | Half B |
59 | 261954 | White | Green | White | N-rectangle | Mostly B |
60 | 261955 | Bronze-green | Bronze-green | Green | N-rectangle | Mostly A |
61 | 261967 | White | White | White | N-rectangle | Mostly B |
62 | 261978 | Red | Red | White | Spherical | Half B |
63 | 261989 | White | White | White | N-rectangle | Mostly B |
64 | 261995 | White | White | White | N-rectangle | Half B |
65 | 262006 | White | White | White | Egg | Mostly B |
66 | 262018 | White | White | White | Rectangle | Mostly B |
67 | 262022 | White | White | White | N-rectangle | Mostly B |
68 | 262023 | White | White | White | W-rectangle | Half B |
69 | 262031 | Red | Red | White | Spherical | Half B |
70 | 262032 | Bronze-green | Bronze-green | Green | W-rectangle | Above |
71 | 262036 | White | Green | Red | Spherical | Half B |
72 | 262037 | White | White | White | N-rectangle | Mostly B |
73 | 262044 | White | White | White | Almond | Half B |
74 | 262049 | White | Green | Red | Spherical | Half B |
75 | 262050 | Red | Red | White | W-Oval | Half B |
76 | 262057 | White | Green | White | Oval | Mostly B |
77 | 262070 | Red | Red | White | W-Oval | Half B |
78 | 262075 | White | White | White | Spherical | Half B |
79 | 262076 | White | White | White | Spherical | Mostly B |
80 | 264178 | White | Green | Red | Spherical | Half B |
81 | 264180 | White | White | White | Spherical | Mostly B |
82 | 276165 | White | White | White | N-rectangle | Mostly A |
83 | 278269 | White | White | White | W-rectangle | Mostly B |
84 | 278682 | White | Green | Red | W-rectangle | Half B |
85 | 278683 | Bronze-green | Bronze-green | Green | N-rectangle | Mostly A |
86 | 278684 | White | White | White | N-rectangle | Mostly A |
87 | 278685 | White | White | White | N-rectangle | Mostly B |
88 | 278727 | Red | Red | White | Spherical | Mostly A |
89 | 283305 | White | White | White | Spherical | Mostly B |
90 | 283312 | White | Green | White | N-rectangle | Mostly B |
91 | 283317 | Red | Red | White | Spherical | Half B |
92 | 289244 | White | Green | Red | Spherical | Half B |
93 | 291383 | Red | Red | White | Egg | Half B |
94 | 291423 | White | Green | White | Oval | Mostly B |
95 | 291541 | White | White | White | Rectangle | Mostly B |
96 | 293006 | White | White | White | Spherical | Mostly B |
97 | 293008 | White | Green | White | N-rectangle | Mostly B |
98 | 293028 | White | Green | White | Egg | Mostly B |
99 | 293085 | White | Green | White | Oval | Mostly B |
100 | 297120 | White | Green | Red | Spherical | Half B |
101 | 297172 | Red | Red | White | Spherical | Mostly A |
102 | 297174 | White | White | White | Rectangle | Mostly B |
103 | 299326 | Bronze-green | Bronze-green | Green | N-rectangle | Mostly A |
104 | 299453 | Bronze-green | Bronze-green | Green | N-rectangle | Mostly A |
105 | 305085 | White | Green | White | Rectangle | Half B |
106 | 305381 | Bronze | Bronze-green | Green | Rectangle | Mostly A |
107 | 306869 | White | White | White | W-Oval | Mostly B |
108 | 308359 | Red | Red | White | Spherical | Half B |
109 | 308367 | Bronze-green | Bronze-green | Green | Rectangle | Mostly A |
110 | 308418 | White | White | White | N-rectangle | Mostly B |
Con1 | Gwailmu | White | Green | Red | W-rectangle | Half B |
Con2 | Meosjinmaskkalmu | White | Green | White | Oval | Half B |
Con3 | Taecheong | White | Green | White | Oval | Mostly A |
Con4 | Cheong-unmu | White | Green | White | Oval | Half B |
Con5 | Chorongmu | White | Green | White | Gourd | Mostly B |
Con6 | Mansa-hyeongtongmu | White | Green | White | Oval | Half B |
Con7 | Togwanggoldeumu | White | Green | White | Egg | Mostly B |
Con8 | Baeksinaltari | White | Green | White | Gourd | Mostly B |
Con9 | Syupeogiljomu | White | Green | White | Oval | Half B |
Con10 | Seohogoldeumu | White | Green | White | Oval | Half B |
Appendix 2.
Appendix 3.
No. | IT No. |
Glucoraphasatin (Average ± SDz) |
Glucoraphenin (Average ± SDz) |
Glucobrassicin (Average ± SDz) |
Glucoerucin (Average ± SDz) |
Glucoberteroin (Average ± SDz) |
Total glucosinolate (Average ± SDz) |
DPPH activity (%) |
1 | 32729 |
11179.03 ± 925.59 |
2656.67 ± 263.93 |
124.37 ± 11.06 |
319.76 ± 34.11 |
24.41 ± 1.81 |
14304.24 ± 1234.00 |
22.29 ± 0.50 |
2 | 100590 |
10939.58 ± 1419.28 |
1778.49 ± 326.62 |
134.18 ± 19.91 |
360.22 ± 55.2 |
35.83 ± 1.74 |
13248.31 ± 1814.64 |
27.60 ± 1.46 |
3 | 100649 |
2793.50 ± 266.26 |
546.68 ± 66.83 |
28.93 ± 3.83 |
27.23 ± 2.01 |
4.36 ± 0.69 |
3400.70 ± 338.45 |
26.02 ± 0.90 |
4 | 100678 |
984.87 ± 121.39 |
472.46 ± 58.10 |
8.27 ± 1.37 |
9.50 ± 1.06 |
0.66 ± 0.32 |
1475.75 ± 181.93 |
19.08 ± 1.52 |
5 | 100684 |
5945.12 ± 390.38 |
1713.83 ± 149.23 |
220.70 ± 19.95 |
133.37 ± 13.98 |
7.90 ± 0.49 |
8020.92 ± 571.75 |
30.90 ± 1.76 |
6 | 100689 |
975.52 ± 126.22 |
747.44 ± 91.26 |
9.40 ± 1.31 |
13.62 ± 1.78 |
1.40 ± 0.52 |
1747.39 ± 220.09 |
31.48 ± 0.68 |
7 | 100691 |
6291.06 ± 689.13 |
1917.39 ± 198.31 |
39.06 ± 5.74 |
102.17 ± 11.39 |
7.99 ± 1.19 |
8357.66 ± 900.7 |
31.26 ± 0.79 |
8 | 100695 |
8476.9 ± 735.11 |
2411.44 ± 167.18 |
288.21 ± 37.26 |
349.47 ± 23.76 |
34.62 ± 2.69 |
11560.65 ± 964.07 |
20.80 ± 0.49 |
9 | 102376 |
11261.78 ± 1145.72 |
1502.84 ± 220.68 |
76.77 ± 8.16 |
324.42 ± 40.68 |
14.46 ± 1.53 |
13180.27 ± 1411.31 |
15.70 ± 1.29 |
10 | 102378 |
11695.28 ± 1055.43 |
2455.14 ± 398.31 |
312.01 ± 23.76 |
291.55 ± 36.15 |
20.95 ± 1.55 |
14774.93 ± 1514.06 |
14.70 ± 2.24 |
11 | 102395 |
5475.85 ± 383.85 |
1328.9 ± 162.74 |
96.22 ± 12.31 |
124.27 ± 7.75 |
7.59 ± 1.52 |
7032.84 ± 564.16 |
21.60 ± 0.21 |
12 | 102560 |
9565.98 ± 355.89 |
2899.40 ± 124.44 |
184.77 ± 5.41 |
378.23 ± 9.46 |
15.62 ± 0.28 |
13043.99 ± 492.01 |
16.02 ± 2.06 |
13 | 103802 |
2815.52 ± 219.04 |
920.06 ± 104.04 |
94.09 ± 10.92 |
36.37 ± 2.68 |
4.59 ± 0.54 |
3870.64 ± 336.22 |
19.18 ± 0.42 |
14 | 103811 |
210.71 ± 18.73 |
57.83 ± 9.42 |
0.87 ± 0.24 |
4.72 ± 0.09 |
0.24 ± 0.15 |
274.37 ± 23.51 |
10.43 ± 2.22 |
15 | 104055 |
2462.68 ± 284.12 |
1045.55 ± 115.08 |
55.75 ± 7.52 |
36.83 ± 4.43 |
3.47 ± 0.86 |
3604.28 ± 411.71 |
18.29 ± 2.17 |
16 | 112253 |
252.61 ± 26.96 |
43.70 ± 5.47 |
3.93 ± 0.31 |
3.70 ± 0.65 |
0.18 ± 0.09 |
304.12 ± 32.98 |
12.47 ± 0.98 |
17 | 112255 |
7013.84 ± 811.76 |
1871.35 ± 326.49 |
122.35 ± 12.8 |
98.84 ± 13.09 |
11.33 ± 0.69 |
9117.71 ± 1161.6 |
16.09 ± 2.80 |
18 | 112257 |
9441.1 ± 750.17 |
936.32 ± 120.22 |
82.20 ± 6.39 |
215.41 ± 21.63 |
11.19 ± 0.72 |
10686.21 ± 897.43 |
19.89 ± 2.80 |
19 | 112258 |
8996.09 ± 747.33 |
1596.7 ± 212.78 |
33.80 ± 2.64 |
445.24 ± 43.83 |
20.56 ± 1.20 |
11092.39 ± 1006.74 |
18.59 ± 2.03 |
20 | 119000 |
9305.38 ± 986.46 |
2231.75 ± 405.44 |
60.17 ± 8.26 |
284.83 ± 45.92 |
19.36 ± 3.54 |
11901.48 ± 1449.54 |
16.47 ± 1.35 |
21 | 136485 |
8551.15 ± 740.75 |
735.01 ± 75.20 |
43.17 ± 5.30 |
247.89 ± 23.55 |
23.8 ± 1.33 |
9601.02 ± 830.78 |
32.49 ± 2.57 |
22 | 136498 |
3116.76 ± 306.78 |
901.86 ± 105.92 |
29.20 ± 2.68 |
48.98 ± 5.56 |
2.89 ± 0.17 |
4099.69 ± 417.29 |
52.69 ± 0.59 |
23 | 166993 |
508.27 ± 41.06 |
379.03 ± 34.81 |
4.71 ± 0.18 |
6.54 ± 0.97 |
0.38 ± 0.25 |
898.94 ± 77.13 |
18.29 ± 0.20 |
24 | 166995 |
5828.08 ± 876.35 |
1862.04 ± 430.71 |
225.72 ± 32.85 |
191.62 ± 39.33 |
4.67 ± 0.37 |
8112.13 ± 1377.45 |
28.77 ± 1.54 |
25 | 166997 |
423.77 ± 49.42 |
207.6 ± 27.75 |
2.08 ± 0.33 |
6.37 ± 0.39 |
0.29 ± 0.26 |
640.11 ± 77.01 |
21.26 ± 0.30 |
26 | 185738 |
691.98 ± 50.89 |
316.58 ± 44.00 |
8.12 ± 0.19 |
16.64 ± 2.18 |
0.49 ± 0.29 |
1033.8 ± 96.62 |
18.72 ± 1.10 |
27 | 188102 |
1615.38 ± 212.87 |
924.99 ± 132.93 |
18.92 ± 2.07 |
23.6 ± 4.24 |
2.21 ± 0.28 |
2585.11 ± 349.14 |
16.35 ± 1.66 |
28 | 203316 |
3715.43 ± 452.50 |
1964.29 ± 284.38 |
158.21 ± 17.09 |
74 ± 12.10 |
9.11 ± 1.91 |
5921.04 ± 766.50 |
20.27 ± 1.55 |
29 | 203531 |
3095.72 ± 309.97 |
960.01 ± 100.11 |
6.43 ± 0.65 |
51.49 ± 7.02 |
4.38 ± 0.48 |
4118.03 ± 418.05 |
16.3 ± 1.14 |
30 | 204160 |
4668.71 ± 590.22 |
1118.75 ± 181.58 |
59.03 ± 8.37 |
98.51 ± 14.91 |
9.54 ± 1.09 |
5954.54 ± 795 |
32.84 ± 0.63 |
31 | 208400 |
814.68 ± 102.08 |
499.81 ± 132.81 |
5.28 ± 0.80 |
13.73 ± 1.72 |
0.49 ± 0.52 |
1334 ± 237.12 |
59.81 ± 0.69 |
32 | 209937 |
3424.68 ± 322.98 |
1808.17 ± 184.18 |
88.7 ± 8.17 |
62.04 ± 7.73 |
8.31 ± 1.33 |
5391.9 ± 520.24 |
23.78 ± 0.64 |
33 | 209974 |
5093.86 ± 688.86 |
1111.8 ± 163.98 |
90.38 ± 12.64 |
172.77 ± 33.28 |
5.36 ± 0.62 |
6474.17 ± 890.91 |
27.58 ± 1.73 |
34 | 210203 |
4050.15 ± 447.00 |
814.92 ± 85.05 |
8.69 ± 1.15 |
89.41 ± 9.81 |
13.08 ± 1.31 |
4976.25 ± 537.67 |
24.90 ± 0.34 |
35 | 213153 |
662.76 ± 62.28 |
510.75 ± 42.22 |
1.56 ± 0.04 |
12.63 ± 1.68 |
0.41 ± 0.11 |
1188.11 ± 105.84 |
18.81 ± 0.51 |
36 | 215011 |
12921.97 ± 1252.18 |
2527.07 ± 295.34 |
134.99 ± 15.18 |
440.03 ± 37.63 |
49.69 ± 5.37 |
16073.75 ± 1549.92 |
28.79 ± 2.41 |
37 | 215079 |
9803.19 ± 806.83 |
1700.19 ± 165.18 |
26.87 ± 1.41 |
348.51 ± 36.71 |
19.78 ± 1.74 |
11898.55 ± 1004.35 |
12.85 ± 1.84 |
38 | 218925 |
11321.86 ± 1208.50 |
2160.83 ± 292.20 |
165.04 ± 11.76 |
512.00 ± 66.73 |
18.62 ± 3.07 |
14178.35 ± 1579.41 |
26.19 ± 0.88 |
39 | 220675 |
7562.46 ± 1027.86 |
3389.28 ± 570.44 |
296.17 ± 45.93 |
199.11 ± 36.77 |
23.76 ± 2.53 |
11470.77 ± 1667.16 |
19.78 ± 1.69 |
40 | 221952 |
8634.13 ± 975.11 |
3140.49 ± 364.04 |
338.87 ± 36.89 |
305.60 ± 30.6 |
43.56 ± 3.06 |
12462.65 ± 1402.28 |
30.83 ± 2.51 |
41 | 221955 |
7544.73 ± 930.51 |
3883.83 ± 522.85 |
28.75 ± 6.21 |
249.00 ± 28.87 |
16.74 ± 1.17 |
11723.95 ± 1461.62 |
33.26 ± 1.74 |
42 | 221956 |
7179.87 ± 775.79 |
2988.78 ± 407.06 |
84.45 ± 5.87 |
227.86 ± 31.48 |
17.77 ± 0.82 |
10498.74 ± 1200.09 |
21.33 ± 1.51 |
43 | 221958 |
325.33 ± 36.35 |
174.68 ± 19.07 |
5.33 ± 0.55 |
6.82 ± 1.01 |
0.15 ± 0.16 |
512.31 ± 56.99 |
18.79 ± 0.36 |
44 | 221959 |
1407.71 ± 166.67 |
616.06 ± 115.42 |
23.68 ± 2.93 |
27.61 ± 2.34 |
1.21 ± 0.43 |
2076.27 ± 259.33 |
34.57 ± 0.56 |
45 | 223576 |
9678.29 ± 1323.37 |
4284.93 ± 551.12 |
224.54 ± 25.15 |
319.74 ± 63.01 |
27.68 ± 6.77 |
14535.19 ± 1926.59 |
19.13 ± 1.24 |
46 | 228857 |
8046.57 ± 861.63 |
3748.14 ± 391.37 |
203.22 ± 31.25 |
291.04 ± 31.17 |
29.59 ± 3.50 |
12318.56 ± 1309.22 |
30.39 ± 1.30 |
47 | 228870 |
11638.83 ± 1215.04 |
5105.15 ± 531.94 |
631.39 ± 129.13 |
371.71 ± 66.63 |
42.26 ± 5.75 |
17789.35 ± 1943.58 |
15.50 ± 1.49 |
48 | 250738 |
10414.1 ± 1215.68 |
1451.88 ± 179.06 |
113.52 ± 13.35 |
319.07 ± 56.43 |
23.71 ± 3.37 |
12322.28 ± 1465.38 |
20.90 ± 2.22 |
49 | 250765 |
5443.16 ± 507.96 |
1471.51 ± 169.37 |
33.07 ± 3.07 |
158.79 ± 22.78 |
6.64 ± 0.84 |
7113.17 ± 702.02 |
17.24 ± 2.05 |
50 | 250775 |
6514.06 ± 588.64 |
2636.95 ± 282.67 |
76.41 ± 13.88 |
187.94 ± 17.38 |
20.89 ± 1.41 |
9436.24 ± 902.71 |
24.64 ± 1.09 |
51 | 250777 |
1161.23 ± 62.82 |
791.86 ± 133.05 |
35.20 ± 0.57 |
20.95 ± 0.95 |
1.62 ± 0.13 |
2010.85 ± 190.33 |
17.84 ± 2.31 |
52 | 250788 |
5719.88 ± 455.08 |
3293.35 ± 115.39 |
99.63 ± 6.72 |
174.70 ± 22.87 |
9.98 ± 0.35 |
9297.54 ± 599.10 |
23.80 ± 0.88 |
53 | 250790 |
7527.83 ± 836.51 |
4421.21 ± 418.65 |
129.48 ± 12.02 |
192.34 ± 35.84 |
13.09 ± 2.04 |
12283.96 ± 1257.89 |
40.63 ± 12.15 |
54 | 250792 |
6315.22 ± 618.27 |
1035.81 ± 107.50 |
246.75 ± 21.10 |
138.07 ± 20.15 |
13.6 ± 0.83 |
7749.45 ± 759.18 |
16.00 ± 3.17 |
55 | 250794 |
7707.88 ± 564.45 |
2828.78 ± 381.80 |
186.71 ± 24.00 |
214.64 ± 23.35 |
23.55 ± 1.12 |
10961.55 ± 982.87 |
22.08 ± 2.48 |
56 | 261944 |
4715.3 ± 607.44 |
496.67 ± 64.25 |
119.13 ± 15.15 |
100.22 ± 11.55 |
5.38 ± 0.39 |
5436.7 ± 697.13 |
28.00 ± 0.97 |
57 | 261947 |
8035.35 ± 729.44 |
3329.76 ± 180.91 |
70.63 ± 8.33 |
263.42 ± 41.36 |
13.94 ± 3.22 |
11713.1 ± 937.09 |
27.53 ± 3.00 |
58 | 261953 |
4415.55 ± 456.00 |
1237.84 ± 144.58 |
73.36 ± 9.98 |
79.53 ± 7.91 |
7.44 ± 0.99 |
5813.72 ± 613.45 |
32.54 ± 1.56 |
59 | 261954 |
6710.18 ± 713.28 |
2105.59 ± 250.08 |
64.73 ± 12.06 |
126.48 ± 13.04 |
6.60 ± 0.21 |
9013.58 ± 987.87 |
14.61 ± 1.67 |
60 | 261955 |
7185.08 ± 813.15 |
1960.97 ± 255.23 |
196.25 ± 28.85 |
187.04 ± 18.21 |
17.68 ± 1.06 |
9547.02 ± 1115.79 |
21.06 ± 0.80 |
61 | 261967 |
8442.83 ± 1006.26 |
3438.58 ± 446.30 |
124.64 ± 18.12 |
262.36 ± 25.49 |
23.77 ± 1.89 |
12292.18 ± 1482.31 |
21.88 ± 1.94 |
62 | 261978 |
974.68 ± 48.05 |
302.64 ± 22.48 |
7.83 ± 0.38 |
13.93 ± 0.93 |
0.79 ± 0.32 |
1299.86 ± 68.28 |
24.38 ± 3.71 |
63 | 261989 |
10760.66 ± 977.97 |
3121.63 ± 339.26 |
26.72 ± 2.51 |
520.92 ± 77.72 |
32.17 ± 1.59 |
14462.1 ± 1329.74 |
20.40 ± 2.37 |
64 | 261995 |
2167.39 ± 226.85 |
1547.31 ± 160.31 |
2.98 ± 0.19 |
40.01 ± 5.13 |
2.80 ± 0.31 |
3760.5 ± 370.32 |
18.38 ± 3.10 |
65 | 262006 |
423.06 ± 22.30 |
248.22 ± 15.75 |
4.21 ± 0.13 |
7.90 ± 0.20 |
0.26 ± 0.09 |
683.65 ± 30.19 |
21.39 ± 1.64 |
66 | 262018 |
147.42 ± 6.48 |
29.23 ± 1.59 |
0.65 ± 0.07 |
2.58 ± 0.18 |
0.05 ± 0.06 |
179.92 ± 7.33 |
24.59 ± 2.02 |
67 | 262022 |
8475.89 ± 920.77 |
4146.82 ± 408.09 |
38.22 ± 2.66 |
310.78 ± 43.94 |
28.01 ± 2.13 |
12999.72 ± 1372.38 |
24.59 ± 2.17 |
68 | 262023 |
184.02 ± 22.47 |
59.04 ± 8.60 |
1.36 ± 0.13 |
2.40 ± 0.09 |
0.00 ± 0.00 |
246.82 ± 31.22 |
27.57 ± 2.23 |
69 | 262031 |
1082.49 ± 98.07 |
778.41 ± 50.81 |
10.68 ± 0.77 |
32.74 ± 3.86 |
3.51 ± 1.01 |
1907.83 ± 151.27 |
27.64 ± 3.18 |
70 | 262032 |
772.05 ± 74.89 |
730.38 ± 35.82 |
13.27 ± 1.10 |
15.66 ± 1.79 |
1.40 ± 0.48 |
1532.76 ± 103.77 |
17.43 ± 1.98 |
71 | 262036 |
7756.15 ± 736.74 |
3407.97 ± 228.30 |
533.12 ± 50.83 |
150.55 ± 24.20 |
11.75 ± 1.03 |
11859.54 ± 1018.87 |
45.69 ± 3.02 |
72 | 262037 |
7161.74 ± 776.01 |
2300.09 ± 265.61 |
12.54 ± 1.41 |
202.79 ± 35.52 |
15.63 ± 1.85 |
9692.79 ± 1066.16 |
15.66 ± 0.80 |
73 | 262044 |
694.95 ± 64.48 |
313.93 ± 21.21 |
16.01 ± 1.00 |
9.62 ± 0.98 |
0.20 ± 0.11 |
1034.71 ± 83.71 |
19.63 ± 1.93 |
74 | 262049 |
1747.75 ± 161.08 |
749.16 ± 100.01 |
143.81 ± 16.09 |
17.98 ± 1.41 |
2.16 ± 0.72 |
2660.86 ± 279.13 |
38.75 ± 1.29 |
75 | 262050 |
2529.17 ± 263.58 |
973.73 ± 124.64 |
50.72 ± 6.24 |
51.29 ± 5.31 |
5.07 ± 0.52 |
3609.98 ± 399.91 |
31.08 ± 1.87 |
76 | 262057 |
1587.23 ± 144.12 |
628.61 ± 72.63 |
14.36 ± 1.54 |
24.32 ± 2.05 |
1.89 ± 0.39 |
2256.41 ± 220.03 |
19.37 ± 1.95 |
77 | 262070 |
2899.72 ± 228.78 |
1012.70 ± 48.97 |
106.6 ± 5.25 |
41.77 ± 3.90 |
3.43 ± 0.60 |
4064.22 ± 278.41 |
18.43 ± 5.71 |
78 | 262075 |
4921.87 ± 389.66 |
2323.96 ± 179.26 |
22.73 ± 1.28 |
118.28 ± 11.36 |
9.53 ± 0.73 |
7396.36 ± 555.24 |
23.91 ± 2.95 |
79 | 262076 |
8207.53 ± 662.86 |
3318.56 ± 198.15 |
74.34 ± 4.57 |
276.04 ± 36.80 |
19.23 ± 0.95 |
11895.69 ± 743.41 |
27.9 ± 3.08 |
80 | 264178 |
3420.25 ± 99.52 |
1329.92 ± 135.41 |
148.07 ± 11.80 |
58.97 ± 1.80 |
6.10 ± 0.37 |
4963.3 ± 220.64 |
63.05 ± 33.28 |
81 | 264180 |
4447.28 ± 431.07 |
1930.04 ± 240.28 |
57.13 ± 5.99 |
95.24 ± 7.94 |
11.49 ± 0.78 |
6541.17 ± 684.68 |
28.09 ± 1.21 |
82 | 276165 |
609.89 ± 54.54 |
400.04 ± 36.88 |
10.52 ± 0.49 |
9.58 ± 0.86 |
0.41 ± 0.27 |
1030.44 ± 88.46 |
20.02 ± 4.00 |
83 | 278269 |
3356.76 ± 326.57 |
1093.71 ± 163.91 |
40.52 ± 3.92 |
48.08 ± 4.83 |
2.19 ± 0.28 |
4541.26 ± 494.96 |
18.96 ± 1.68 |
84 | 278682 |
4711.53 ± 382.95 |
1110.76 ± 171.65 |
591.27 ± 42.72 |
64.53 ± 6.23 |
4.90 ± 0.99 |
6482.99 ± 543.26 |
55.17 ± 4.43 |
85 | 278683 |
11663.38 ± 1257.80 |
2894.93 ± 358.63 |
703.41 ± 76.07 |
454.01 ± 80.97 |
40.22 ± 1.09 |
15755.96 ± 1713.63 |
16.07 ± 2.60 |
86 | 278684 |
8767.59 ± 1025.95 |
2648.30 ± 217.99 |
72.76 ± 8.83 |
347.88 ± 63.10 |
18.52 ± 2.29 |
11855.05 ± 1309.80 |
12.68 ± 16.25 |
87 | 278685 |
1316.78 ± 102.33 |
1104.85 ± 72.23 |
5.29 ± 0.38 |
21.07 ± 2.20 |
0.58 ± 0.25 |
2448.57 ± 133.23 |
4.26 ± 25.19 |
88 | 278727 |
1266.86 ± 129.63 |
792.83 ± 41.70 |
18.25 ± 1.19 |
18.19 ± 1.93 |
1.31 ± 0.06 |
2097.44 ± 143.08 |
46.29 ± 0.53 |
89 | 283305 |
12816.85 ± 1468.17 |
2342.25 ± 298.43 |
96.38 ± 9.67 |
466.24 ± 47.87 |
31.53 ± 2.14 |
15753.25 ± 1825.81 |
31.67 ± 1.56 |
90 | 283312 |
4895.8 ± 372.14 |
1234.99 ± 26.07 |
27.01 ± 2.09 |
97.90 ± 12.57 |
4.57 ± 0.37 |
6260.26 ± 389.30 |
17.95 ± 2.52 |
91 | 283317 |
5192.73 ± 526.49 |
1330.03 ± 147.24 |
96.69 ± 9.80 |
102.34 ± 16.28 |
9.82 ± 0.96 |
6731.6 ± 678.03 |
14.21 ± 1.99 |
92 | 289244 |
5002.57 ± 662.49 |
1517.41 ± 255.09 |
324.15 ± 50.58 |
82.35 ± 10.63 |
5.89 ± 1.64 |
6932.37 ± 979.65 |
44.99 ± 1.48 |
93 | 291383 |
1523.63 ± 196.73 |
445.56 ± 63.93 |
14.41 ± 1.47 |
28.15 ± 2.76 |
2.65 ± 0.25 |
2014.41 ± 263.61 |
36.14 ± 0.46 |
94 | 291423 |
9723.58 ± 1019.92 |
1730.22 ± 246.64 |
334.58 ± 42.10 |
466.17 ± 36.92 |
30.45 ± 2.61 |
12285.01 ± 1346.28 |
14.88 ± 0.14 |
95 | 291541 |
1913.56 ± 76.47 |
976.86 ± 113.15 |
8.10 ± 0.22 |
39.47 ± 1.05 |
2.20 ± 0.19 |
2940.19 ± 187.07 |
21.94 ± 1.73 |
96 | 293006 |
7856.62 ± 573.90 |
1144.05 ± 52.40 |
28.61 ± 2.51 |
251.52 ± 26.31 |
13.26 ± 0.79 |
9294.06 ± 625.16 |
17.01 ± 2.99 |
97 | 293008 |
6179.58 ± 630.56 |
1514.71 ± 91.55 |
14.66 ± 1.52 |
161.65 ± 23.71 |
12.20 ± 2.30 |
7882.8 ± 745.08 |
22.57 ± 1.86 |
98 | 293028 |
8425.24 ± 679.54 |
934.71 ± 75.56 |
32.62 ± 2.15 |
249.98 ± 31.27 |
13.41 ± 1.70 |
9655.96 ± 747.74 |
20.73 ± 2.25 |
99 | 293085 |
7402.03 ± 706.58 |
2230.55 ± 229.32 |
93.13 ± 23.58 |
237.61 ± 22.52 |
14.25 ± 1.36 |
9977.56 ± 976.05 |
24.63 ± 0.41 |
100 | 297120 |
6933.66 ± 428.54 |
2029.14 ± 106.05 |
449.10 ± 30.50 |
170.32 ± 16.06 |
12.56 ± 0.18 |
9594.78 ± 504.88 |
46.39 ± 0.74 |
101 | 297172 |
469.18 ± 44.07 |
163.82 ± 11.35 |
2.03 ± 0.14 |
6.77 ± 0.70 |
0.19 ± 0.07 |
641.98 ± 52.73 |
30.53 ± 3.13 |
102 | 297174 |
9182.81 ± 914.73 |
2213.60 ± 213.38 |
203.22 ± 20.46 |
320.68 ± 22.55 |
20.87 ± 1.01 |
11941.19 ± 1162.16 |
18.98 ± 0.05 |
103 | 299326 |
9389.02 ± 492.80 |
3128.66 ± 149.46 |
380.53 ± 24.92 |
363.06 ± 34.87 |
43.72 ± 4.16 |
13305 ± 612.02 |
29.43 ± 3.52 |
104 | 299453 |
12495.36 ± 805.01 |
4946.47 ± 302.76 |
403.09 ± 17.58 |
534.53 ± 51.15 |
59.07 ± 3.24 |
18438.52 ± 970.31 |
17.77 ± 3.55 |
105 | 305085 |
6097.55 ± 425.32 |
2012.43 ± 326.01 |
255.72 ± 33.40 |
154.40 ± 12.75 |
11.19 ± 0.80 |
8531.3 ± 792.97 |
21.49 ± 0.56 |
106 | 305381 |
5909.04 ± 581.05 |
1970.40 ± 280.71 |
372.52 ± 52.71 |
212.09 ± 26.37 |
20.16 ± 4.22 |
8484.21 ± 941.19 |
28.34 ± 0.29 |
107 | 306869 |
10571.92 ± 1141.35 |
5653.20 ± 731.55 |
1243.88 ± 187.47 |
514.58 ± 49.20 |
34.30 ± 2.97 |
18017.88 ± 2108.07 |
29.77 ± 0.94 |
108 | 308359 |
7635.26 ± 195.76 |
3196.07 ± 309.54 |
166.23 ± 3.77 |
218.03 ± 9.36 |
16.94 ± 0.74 |
11232.53 ± 421.18 |
21.61 ± 2.78 |
109 | 308367 |
5906.01 ± 157.17 |
1353.42 ± 168.61 |
250.59 ± 11.37 |
198.84 ± 7.28 |
27.66 ± 2.92 |
7736.50 ± 246.11 |
22.85 ± 1.54 |
110 | 308418 |
123.76 ± 16.36 |
26.28 ± 3.50 |
1.27 ± 0.11 |
2.97 ± 0.20 |
0.02 ± 0.02 |
154.30 ± 20.09 |
29.52 ± 0.16 |
Con1 | Gwailmu |
2531.25 ± 296.06 |
1247.67 ± 191.43 |
268.36 ± 44.46 |
40.86 ± 4.88 |
5.31 ± 1.24 |
4093.45 ± 536.93 |
43.31 ± 0.54 |
Con2 |
Meosjin maskkalmu |
4702 ± 522.40 |
694.89 ± 101.45 |
30.79 ± 4.29 |
112.71 ± 11.91 |
4.72 ± 1.30 |
5545.11 ± 641.26 |
26.91 ± 1.04 |
Con3 | Taecheong |
6437.09 ± 545.29 |
509.63 ± 51.17 |
7.94 ± 1.22 |
120.97 ± 16.81 |
5.05 ± 0.93 |
7080.67 ± 610.67 |
23.36 ± 1.83 |
Con4 |
Cheong- unmu |
6210.01 ± 775.16 |
898.02 ± 122.53 |
27.43 ± 4.23 |
146.01 ± 17.69 |
10.01 ± 1.78 |
7291.48 ± 920.80 |
21.66 ± 2.47 |
Con5 | Chorongmu |
6781.61 ± 611.26 |
532.23 ± 62.75 |
25.82 ± 3.70 |
123.83 ± 8.55 |
8.89 ± 0.35 |
7472.38 ± 685.75 |
30.67 ± 0.49 |
Con6 |
Mansa- hyeong tongmu |
7019.84 ± 922.97 |
984.58 ± 148.58 |
44.71 ± 7.02 |
189.20 ± 26.21 |
12.25 ± 1.57 |
8250.57 ± 1105.98 |
26.58 ± 0.92 |
Con7 |
Togwang goldeumu |
7968.49 ± 453.95 |
991.70 ± 64.54 |
37.75 ± 2.59 |
169.24 ± 16.43 |
8.83 ± 0.85 |
9176.01 ± 492.2 |
21.92 ± 3.48 |
Con8 |
Baeksi naltari |
7989.39 ± 755.68 |
1144.97 ± 126.38 |
185.86 ± 32.79 |
146.49 ± 14.07 |
10.29 ± 1.77 |
9477.00 ± 929.50 |
26.91 ± 0.43 |
Con9 |
Syupeo giljomu |
9053.54 ± 738.81 |
1307.13 ± 136.26 |
48.53 ± 6.08 |
212.05 ± 17.12 |
15.36 ± 1.48 |
10636.61 ± 894.57 |
24.51 ± 0.20 |
Con10 |
Seoho goldeumu |
9247.03 ± 706.53 |
1256.86 ± 111.22 |
59.03 ± 8.46 |
292.94 ± 21.42 |
17.25 ± 1.53 |
10873.12 ± 843.35 |
29.27 ± 0.44 |