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HORTSCIENCE 53(12):1732–1736. 2018. https://doi.org/10.21273/HORTSCI13448-18 complete , but also soy food helps to reduce health risks such as cardiovascular disease, osteoporosis, and cancers (Messina, Evaluation of Lines for 1999; Messina and Wu, 2009). Edamame consumption improves human health by Edamame Yield Traits and Trait providing high levels of protein and all es- sential amino acids in well-balanced profiles, Genetic Correlation along with other nutritional compounds like isoflavones. Guo-Liang Jiang1, Laban K. Rutto, and Shuxin Ren The food-type soybean industry has spent Agricultural Research Station, Virginia State University, P.O. Box 9061, more than 20 years growing from door-to- Petersburg, VA 23806 door sales to an integrated supply chain throughout the United States (Soyfoods As- Additional index words. vegetable soybean, Glycine max, yield, heritability, genetic sociation of North America, 2011). The correlation market demand for vegetable has significantly increased. In Arkansas, eda- Abstract Glycine max . Edamame is a vegetable or specialty soybean ( (L.) Merr.) with high mame is commercially grown and an nutrition and market value. The market demand for edamame has significantly edamame-processing plant is in operation increased in the United States since its health and nutritional benefits became recognized. (McBryde, 2012). Edamame (fresh or frozen However, there are a limited number of domestically developed or improved edamame pods and/or seeds) is now available in super- cultivars in the United States, and the knowledge of edamame is very limited. In this markets (e.g., Walmart), grocery stores (e.g., study, 86 breeding lines and cultivars of maturity group (MG) V and VI developed in the Kroger), wholesale stores (Sam’s Club and United States were evaluated in replicated field trials for edamame yield and agronomic Costco), farmers markets, and/or restaurants. traits in Virginia in 2015 and 2016. The results indicated that there were significant The increased demand has resulted in a steady differences among the genotypes and between years in all the traits investigated (plant increase in land acreage under vegetable height, fresh biomass, pod yield, pod ratio, fresh seed yield, seed ratio, and 100-seed soybeans (Binder, 2010). Edamame provides weights), but the yearly differences for dried 100-seed weight and dried-to-fresh ratio of a good option of crop farming, especially for seeds were insignificant. Genotype-by-year interaction effects were not significant in small-scale farmers and urban agriculture most cases. Estimates of the broad sense heritability varied with traits, from 23% to 88%. growers who want to increase income by Coefficients of phenotypic and genotypic correlation were mostly low, but fresh pod and growing such a high-value niche crop. How- seed yields were highly correlated. Fresh biomass exhibited a positive phenotypic ever, edamame products sold in markets correlation with pod and seed yields, but the genotypic correlation coefficients were across the United States are mainly imported not significant. Twelve breeding lines were preliminarily identified to have greater from China and other Asian countries or edamame yield and desired traits. The information generated in this study will be helpful regions and are mostly marketed frozen. One for edamame breeding and commercial production. of the important reasons for this status is that there is a lack of domestically developed or improved edamame cultivars in the United Soybean [Glycine max (L.) Merr.] is an only to corn in terms of total acreage and States and also lack of affordable seed. There- important crop grown worldwide for the economic value. However, vegetable soybean fore, growing edamame faces challenges and/ provision of vegetable oil for human con- is relatively new to North America, although it or problems. From the point of plant breeding, sumption and protein meal for animal feeds. was once grown in the United States during the major problems include limited genetic Edamame, a Japanese term, is a type of WWII, compared with the history of edamame resources, lodging, inferior plant structure or specialty soybeans, also called vegetable grown in Asia for many centuries (Shurtleff type, susceptibility to seed diseases, low yield soybean and maodou in China. In the United and Aoyagi, 2009). Edamame is harvested at potential, and shattering, which is important States, soybean is a major field crop, second the R6 growth stage, when the pods and seeds for edamame seed production. are still green (Fehr et al., 1971), different Correspondingly, research on edamame, from general-purpose soybeans, which are in particular plant breeding and genetics harvested after full maturity (R8 stage). studies, has rarely been reported in the United Received for publication 6 Aug. 2018. Accepted Comparatively speaking, edamame is for publication 17 Sept. 2018. States. Rao et al. (2002) analyzed fresh green grown on a much smaller scale but has pod and seed yields in 12 vegetable or large- This study was supported in part by U.S. De- greater market and nutrition values than partment of Agriculture-National Institute of Food seeded soybean cultivars/genotypes from and Agriculture Evans-Allen Research Program general-purpose soybean (Liu, 1999). It has Japan and China and two adapted U.S. and USDA-NIFA Capacity Building Grant (CBG) been reported that the net return per acre for cultivars. Zhang and Kyei-Boahen (2007) Program (funding awarded to G.-L. J.). growing edamame was $259 for the whole- evaluated five traits including fresh pod We are grateful to the Virginia State University sale fresh market in Kentucky (Ernst and weight in 23 edamame cultivars from MG students Haley Berry, Rameka Tylor, and Zhane Woods, 2001), $600 to $750 for farmers’ III to VII in Mississippi. Mebrahtu and Slade for their assistance on the projects. We also markets in Ohio (Bernick, 2009), and overall Mohamed (2006) analyzed genetic variation especially thank Ronald Bowen, former Plant ranged from $400 to $1300 in the United for green pod yield and quality in 31 vegeta- Science Manager retired from Virginia State Uni- States (Binder, 2010) compared with $350 to versity, for his technical support and help in field ble soybean genotypes from MG III to VI. trials and data collection, and James B. Holland, $600 of the gross income per acre for growing More recently, Ogles et al. (2016) evaluated USDA-ARS Plant Science Research Unit and De- general-purpose soybean. Soy food provides 11 selected edamame cultivars of four MGs partment of Crop and Soil Sciences at North complete protein with all essential amino for adaptability and production in central Carolina State University, for his advice on the acids and benefits human health (Messina, Alabama. By comparing 136 entries from SAS program in analysis of genotypic correlation. 1999). Although soybean historically has not 22 resources with 14 grain-type cultivars This article is a contribution of the Virginia State been viewed as an edible crop in the United representing a range of MGs, Williams University, Agricultural Research Station (Journal States, more people have become aware of soy (2015) characterized the vegetable soybean Series No. 352). foods like and edamame, and more people 1Corresponding author. E-mail: [email protected] or germplasm lines for commercial production. [email protected]. are adopting plant-based diets (CBS News, In these studies, differential maturities were This is an open access article distributed under the 2013; Kelley and Sanchez, 2005). The in- involved and the materials mostly originated CC BY-NC-ND license (https://creativecommons. crease of consumption of soy food is not only from out of the United States. To some org/licenses/by-nc-nd/4.0/). because soybean is an important source of extent, the results might be affected by the

1732 HORTSCIENCE VOL. 53(12) DECEMBER 2018 | BREEDING,CULTIVARS,ROOTSTOCKS, AND GERMPLASM RESOURCES

poor adaptation. Use of adapted genotypes narrow harvest-window of edamame, 86 traits i and j, respectively, spi and sgi are with similar maturities in research would re- genotypes of MGs V and VI, selected based the square root of phenotypic and genotypic duce such impacts and provide more useful on similar seed/pod development status (R6 variance for trait i, and spj and sgj are the information for production. Since the 1990s, stage) at sampling dates after excluding MG square root of phenotypic and genotypic the Virginia State University soybean program IV materials, were sampled within a period of variance for trait j. has been dedicated to the development of 1to2weeksfortworeplicationsinbothyears. Genotypic and phenotypic correlation co- edamame cultivars adapted to the United In addition to the four check cultivars described efficients were tested for significance using t States, in particular Virginia and similar envi- previously (‘Asmara’, ‘Mooncake’, ‘N6202-8’, test as t = r/SEr suggested by Robertson ronments (Mebrahtu et al., 2005). The objec- and ‘Randolph’), these selected genotypes in- (1959) and Sharma (1988). tive of this study was to evaluate the fresh cluded 82 F4-derived lines mostly from 18 of edamame yield and related agronomic traits in the diallel crosses made using 10 cultivars and Results and Discussion locally bred soybean cultivars and/or breeding germplasm lines (‘Kanrich’, ‘Late Giant’, lines of MGs V and VI and to analyze the trait ‘Pella’, ‘Tomahomare’, ‘Verde’, PI 379621, Performance of traits. Averages of the correlations, to help the identification and de- PI 399055, PI 506852, V81-1603, and VS95- traits evaluated across 2 years are shown in velopment of adapted superior edamame culti- 50), which were chosen on the basis of the fresh Table 1. There was a very large difference vars with desired traits for the United States. pod yield, nutritional values, and panel taste between the extremes for fresh biomass, pod, test (Mebrahtu and Devine, 2009). Three to and seed yield. Similarly, Zhang and Kyei- Materials and Methods eight lines from each of the 18 crosses were Boahen (2007) also reported a large differ- evaluated in this study. Therefore, the sub- ence from 1,612 to 21,429 kg·ha–1 in fresh Plant materials and field experiments. In sequent analyses were only based on the 86 pod yield among varieties. Frequency distri- total, 152 entries/genotypes of MG IV, V, and genotypes and two replications for 2 years. butions for the traits were similar to a normal VI of soybean were grown in the yield trials Sampling and measurements. At R6 stage distribution (Fig. 1). The CVs indicated that, at the Virginia State University Randolph or the ‘‘green bean’’ stage, all plants within comparatively, the edamame yield of either Research Farm in Ettrick, VA, in 2015 and 1.5 m of an inside row per plot were cut from fresh pods or seeds and biomass showed 2016. Most of the genotypes were the ad- the bottom of their stems and tied together. a larger relative variation, followed by pod vanced lines developed by Virginia State The bundles of samples were weighed on ratio and 100-seed weights, whereas seed University soybean/edamame program and a scale for fresh biomass (kilograms). The ratio, dried-to-fresh seed ratio, and plant derived from different crosses, in addition to sampled plants were then threshed using height exhibited smaller relative variations check cultivars and lines, ‘Asmara’, ‘Moon- a stationary edamame thresher (Mitsuwa & (Table 1). cake’, ‘N6202-8’, and ‘Randolph’. The ex- Co., Ltd., Tokyo, Japan). All the fresh pods ANOVA showed that genotypic effects perimental field is located at 3714#34$N were collected and weighed immediately for were significant at P < 0.01 for all traits and 7725#32$W, south of Chesterfield fresh pod weight (kilograms) and the pro- (Table 1) except dried-to-fresh seed ratio (P < County in Virginia, with a type of sandy soil portion of pods or pod ratio (%). Then, the 0.05). A significant difference between years (series – Bourne, and family – fine, silty pods were stored in a freezer for 4 to 6 weeks also was found in most of the traits, whereas mixed thermic). In late April and/or early for further use. A random subsample of about there was no significant genotype-by-year May, the herbicide Round-Up (Monsanto 100 pods was taken to determine the weight interaction except for plant height, bio- Company, St. Louis, MO) was sprayed to (grams) and seed size (g/100 seed) of the mass, and seed ratio, which was similar to kill weeds in the experimental field, and the fresh beans, as well as the proportion of fresh a previously reported result on pod yield soil was ploughed about 2 weeks before seeds, or seed ratio (%). The subsampled seeds (Carson et al., 2011). Plant height, bio- planting. The field was tilled again immedi- and the remaining pods after subsampling mass, fresh pod and seed yield, and seed ately before planting. were fully dried at 65 to 70 Cfor3weeks ratio in 2015 were greater than those of A randomized complete block design was or so to determine dried 100-seed weight 2016, with a difference of 36.6 cm, 8463.8 –1 –1 –1 used for the experiments with three replicates. (grams). Plant heights (centimeters) were kg·ha , 498.0 kg·ha , 762.4 kg·ha ,and Each genotype was planted in a four-row plot measured from the bottom to the top of the 6.86% on average, respectively (Table 2). with a 4.8-m length and 0.76-m row spacing. main stem, and the average of four measure- Pod ratio and fresh 100-seed weight in For the experiment in 2015, about 23 plants ments per plot was used in analysis. Yield data 2015 were 6.43% and 2.1 g lower than were planted per meter in mid-May of 2015. and biomass were converted into kg·ha–1. those of 2016. For the experiment in 2016, 17 plants per Statistical analysis. Based on a random- Plant height and fresh and dried 100-seed meter were planted in early June of 2016 due ized complete block design, analysis of var- weight exhibited greater estimates of broad to rain and because the population density in iance (ANOVA) was performed using PROC sense heritability than other traits (Table 1). 2015 seemed too high for edamame. The crop GLM in SAS, version 9.4 (SAS Institute Inc., Mebrahtu and Mohamed (2006) reported management was similar to general soybean Cary, NC), and descriptive statistics, mean, a greater heritability for plant height and production, with the herbicides Select (Valent green pod yield as well. However, lower range, and CV also were calculated. Fre- USA Corporation, Walnut Creek, CA) and quency distribution was computed in Micro- estimates of broad sense heritability (<40%) Storm (United Phosphorus, Inc., King of soft Excel 2013 (Redmond, WA). The broad were observed in this study for the edamame Prussia, PA) applied to control grasses and sense heritability was estimated on a geno- yield of both fresh pods and fresh seeds, and broadleaf weeds as needed. The herbicides type mean basis (Fehr, 1987), as the biomass (Table 1). The results indicated were purchased from a local store of the that edamame yield was considerably af- 2 s2=½s2 s2 = s2=ð Þ Southern States Co. and used under an exper- h = g g + gy y+ e r · y fected by environment, but plant height and imental use permit, and the edamame material fresh 100-seed weights were quite stable where s 2 is the genotypic variance, s 2 is harvested was not used for consumption but g gy across years. the genotype-by-year interaction variance, s 2 is research purpose only. e Correlations between years. Although the environmental variance, r is the number The traits investigated included plant edamame yield and related traits varied of replications, and y is the number of years. height (centimeters from the bottom to the greatly with years or environments, coeffi- Coefficients of phenotypic and genotypic top of main stem), fresh biomass of whole cients of Pearson’s correlation of traits be- correlations were computed using SAS codes plants aboveground (kg·ha–1), fresh pod yield tween 2015 and 2016 were statistically described by Holland (2006), as (kg·ha–1), fresh seed yield (kg·ha–1), the significant, except for pod yield (Table 2). s =ðs weight (grams) of fresh and dried 100-seeds, rpij = pij pi·spjÞ; and rgij = sgij=ðsgi·sgjÞ The correlation coefficients averaged 0.412, pod ratio (the percent of pod weight to bio- ranging from 0.163 for pod yield to 0.765 for mass), and seed ratio (the percent of seed to where spij and sgij are the estimated pheno- fresh 100-seed weight. It also was noticed pod weight). In light of limited time for the typic and genotypic covariance between that among the traits, the estimates of broad

HORTSCIENCE VOL. 53(12) DECEMBER 2018 1733 Table 1. Means, ANOVA, and estimates of heritability of edamame traits over 2 years. 2 Trait Mean Range CV (%) MSYr MSGen MSGen/Yr MSe h (%) Plant height (cm) 103.1 ± 10.8 70.7–128.9 10.51 115155** 469.51** 97.58** 41.56 79.22 Biomass (kg·ha–1) 27,259.8 ± 5,016.9 14,645.7–39,556.0 18.40 4710432558** 79528996** 47764627* 34261391 39.94 Pod yield (kg·ha–1) 6,949.8 ± 1,383.8 4,059.3–11,098.0 19.91 20320745* 6061886** 4669339 3660338 22.97 Pod ratio (%) 26.1 ± 3.8 18.3–35.2 14.42 2590.97** 46.44** 24.05 21.60 48.22 Seed ratio (%) 55.8 ± 2.8 49.8–60.8 5.03 3391.41** 27.49** 13.28* 9.52 51.68 Seed yield (kg·ha–1) 3,884.6 ± 830.9 2,241.3–6,050.6 21.39 43799854** 2200494** 1500747 1180166 31.80 Fresh 100-seed weight (g) 39.1 ± 7.0 25.4–63.8 17.82 180.21** 169.12** 20.36 23.16 87.96 Dried 100-seed weight (g) 12.3 ± 2.0 8.4–17.7 16.16 8.52 13.46** 4.02 4.53 70.18 Dried-to-fresh ratio of seeds (%) 31.4 ± 2.2 26.3–36.6 6.90 3.09 15.07* 8.84 9.83 41.35 *, **Significant at P = 0.05 or 0.01, respectively. ANOVA = analysis of variance; MSYr = mean square of years; MSGen = mean square of genotypes; MSGen/Yr = mean square of genotype-by-year interactions; MSe = mean square of errors.

Fig. 1. Frequency distribution of soybean lines for edamame yield and other traits. sense heritability were highly consistent with suppose that the differences and low correla- effect of environmental conditions and lower the Pearson’s correlation coefficients be- tions in edamame yield and biomass between heritability and also due to different density tween years (Tables 1 and 2). We could years were mainly attributed to the larger and planting times.

1734 HORTSCIENCE VOL. 53(12) DECEMBER 2018 Correlations between traits. Pearson’s nation greater than 0.9 for both phenotypic seeds would not necessarily lead to a de- correlations among the traits showed mostly and genotypic correlation (Table 3). This creased edamame yield, although extralarge- similar between the 2 years (data not shown). relationship is obvious because more pods seeded varieties (mature 100-seed weight To better elucidate the relationships between means more seed yield, and thus one trait >25 g) might produce less seed yield for traits, we further computed the phenotypic may be used as a selection indicator for the grain-type soybeans. Selection of fresh 100- and genotypic correlations. As shown in other in breeding. Plant height showed un- seed weight could be practicable and effec- Table 3, most coefficients of phenotypic remarkable or slight correlation with eda- tive in edamame breeding as it had higher and genotypic correlation were low. Of 36 mame yields and seed sizes. Biomass heritability as well. In addition, pod and seed pairs of traits, about two-thirds were signif- exhibited a positive phenotypic correlation ratios were positively correlated, and both icant in phenotypic correlation, but only with fresh pod and seed yield, but their were genetically related to fresh pod and seed six pairs showed a phenotypic correlation genotypic correlations were insignificant. yields. coefficient greater than 0.4, and 13 pairs For mature soybean, the relationship be- Promising lines. Based on the results over exhibited a significant genotypic correla- tween seed size and seed yield has not been years, we preliminarily identified 12 lines tion. Although phenotypic and genotypic conclusively established (Brar and Carter, that may have desired traits (Table 4). These correlations were consistent in many cases, 1993). Interestingly, the genotypic correla- lines exhibited both greater edamame yield the latter was less significant than the former tion between edamame yield and fresh or and 100-seed weight than the averages of all due to larger SEs of genotypic correlation dried 100-seed weight at R6 stage was very genotypes evaluated and were superior to all coefficients. low in this study. The results implied that four check cultivars: ‘Asmara’, ‘Moon Fresh pod and seed yields were highly seed size might not be genetically correlated Cake’, ‘N6202-8’, and ‘Randolph’. On aver- correlated, showing a coefficient of determi- with edamame yields, and selection for large age, the fresh pod and seed yield in these lines

Table 2. Comparison of edamame traits between year means of 2015 and 2016. 2015 2016 Trait Mean Range Mean Range Correlation coefficient Plant height (cm) 121.4 ± 13.7 78.5–147.5 84.8 ± 9.7 62.0–110.4 0.695** Biomass (kg·ha–1) 31,491.7 ± 7,886.9 10,131.2–49,763.8 23,027.9 ± 4,146.7 12,069.1–32,598.4 0.325** Pod yield (kg·ha–1) 7,198.8 ± 2,209.3 1,889.8–12,257.2 6,700.8 ± 1,345.6 3,493.4–9,938.8 0.163 Pod ratio (%) 22.9 ± 4.7 14.6–37.0 29.4 ± 4.6 18.2–44.0 0.319** Seed ratio (%) 59.2 ± 3.5 49.3–67.9 52.3 ± 3.4 44.0–60.2 0.341** Seed yield (kg·ha–1) 4,265.8 ± 1,333.3 1,173.9–7,196.0 3,503.3 ± 729.6 1,779.6–4,957.1 0.232 * Fresh 100-seed weight (g) 38.1 ± 7.5 23.9–62.6 40.2 ± 7.4 24.6–66.3 0.765** Dried 100-seed weight (g) 12.0 ± 2.5 7.0–18.8 12.5 ± 2.1 8.5–18.0 0.530** Dried-to-fresh ratio of seeds (%) 31.4 ± 3.2 22.9–38.7 31.4 ± 2.0 25.7–35.5 0.342** *, **Significant at P = 0.05 or 0.01, respectively.

Table 3. Coefficients of phenotypic (above diagonal) and genotypic (below diagonal) correlations between edamame traits evaluated jointly in 2015 and 2016. Fresh Dried Dried-to-fresh Trait Plant ht Biomass Pod yield Pod ratio Seed ratio Seed yield 100-seed wt 100-seed wt ratio of seeds Plant height 0.275** 0.067 –0.301** –0.090 0.029 –0.135 –0.098 0.064 Biomass 0.563** 0.757** –0.116 –0.237** 0.680** –0.156* –0.145* 0.041 Pod yield 0.224* 0.424 0.518** 0.019 0.971** 0.012 0.091 0.189** Pod ratio –0.425** –0.793** 0.253 0.315** 0.567** 0.194* 0.322** 0.267** Seed ratio –0.347* –0.289 0.683 0.855** 0.244** 0.209** 0.226** 0.060 Seed yield –0.012 0.209 0.950** 0.459* 0.887** 0.060 0.139* 0.198** Fresh 100-seed weight –0.215 –0.255 –0.030 0.173 –0.105 –0.049 0.889** –0.199** Dried 100-seed weight –0.209 –0.258 0.059 0.266 0.019 0.063 0. 978** 0.253** Dried-to-fresh ratio of seeds 0.088 0.077 0.265 0.214 0.571* 0.402 –0.702** –0.553*

*, **Significant at P = 0.05 or 0.01, respectively, according to t test as t = r/SEr.

Table 4. Average of edamame yield and related agronomic traits in the promising soybean lines identified. Biomass Pod yield Seed yield Fresh Dried Dried-to-fresh Genotype Plant ht (cm) (kg·ha–1) (kg·· ha–1) Pod ratio (%) Seed ratio (%) (kg·ha–1) 100-seed wt (g) 100-seed wt (g) ratio of seeds (%) Asmara (CK 1) 94.8 27,631.2 6,935.4 25.2 54.8 3,832.1 38.3 10.8 28.1 Mooncake (CK 2) 103.7 27,369.9 5,844.5 21.4 56.1 3,293.7 35.5 10.7 30.1 N6202-8 (CK 3) 97.1 31,510.6 7,667.1 25.7 53.4 4,151.0 40.2 13.8 34.2 Randolph (CK 4) 98.9 32,909.0 6,410.1 19.3 51.9 3,352.7 28.3 9.2 32.2 VS12-0129 97.8 23,794.8 7,148.3 30.1 58.7 4,222.0 52.4 16.2 30.9 VS12-0128 94.6 27,620.3 7,966.8 29.8 60.2 4,861.8 49.6 14.5 29.3 VS12-0133 99.2 28,337.7 7,708.4 27.8 58.4 4,518.1 49.6 14.6 29.4 VS11-0022 102.1 29,842.5 7,281.3 26.7 51.1 3,738.1 45.2 14.9 33.9 VS12-0176 93.2 25,412.7 7,500.1 30.3 56.3 4,283.8 42.9 13.2 30.7 VS12-0178 91.4 25,739.3 7,955.1 31.8 56.9 4,518.0 43.5 13.6 31.4 VS12-0202 93.0 27,362.2 7,326.8 26.8 57.5 4,201.6 44.2 12.9 29.2 VS12-0108 109.2 31,325.5 8,920.4 29.4 57.3 5,118.6 46.9 15.4 32.6 VS12-0101 109.2 29,285.5 9,251.1 32.4 57.3 5,347.3 46.2 15.2 32.9 VS12-0095 107.8 25,955.1 7,165.1 28.4 58.9 4,314.4 45.8 14.6 32.4 VS12-0094 110.6 29,489.6 8,382.3 28.2 58.2 4,915.4 45.1 13.3 29.8 VS11-0034 109.4 33,432.5 10,711.6 32.2 49.9 5,399.4 42.7 13.6 31.8 Mean of four CKs 98.6 29,855.2 6,714.3 22.9 54.1 3,657.4 35.6 11.1 31.1 LSD0.05 9.0 8,874.4 2,900.7 7.0 4.7 1,647.1 7.3 3.2 4.8 LSD = least significant difference.

HORTSCIENCE VOL. 53(12) DECEMBER 2018 1735 were 7148.3 to 10711.6 kg·ha–1 and 3738.1 to cbsnews.com/news/edamame-is-the-future-of- Mebrahtu, T. and T.E. Devine. 2009. Diallel anal- 5399.4 kg·ha–1, respectively, compared with american-soy-farmers-profits-in-edible-soy-beans- ysis of composition of 10 vegetable the averages of four check cultivars (6714.3 for-people/>. soybean lines. Plant Breed. 128:249–252. and 3657.4 kg·ha–1) (Table 4). The fresh and Ernst, M. and T. Woods. 2001. Marketing challenges Messina, M.J. 1999. Legumes and soybeans: dried 100-seed weights ranged from 42.7 to for emerging crops in Kentucky: Vegetable Overview of their nutritional profiles and soybeans. University of Kentucky Cooperative health effects. Amer. J. Clin. Nutr. 70:439– 52.4 g and 12.9 to 16.2 g, whereas the aver- Extension Publication, Lexington, KY. 16 Feb. 450. ages of the four check cultivars were 35.6 g 2015. . the soy-breast cancer relation. Amer. J. Clin. e.g., VS11-0022, VS12-0176, and VS12-0202, Fehr, W.R. 1987. Principles of cultivar develop- Nutr. 89:1673–1679. also exhibited good resistance to lodging and ment. Vol 1. Theory and technique. Macmillan, Ogles, C.Z., E.A. Guertal, and D.B. Weaver. 2016. shattering, which is important for edamame New York, NY. Edamame cultivar evaluation in Central Ala- seed production (data not shown). It suggested Fehr, W.R., C.E. Caviness, D.T. Burmood, and J.S. bama. Agron. J. 108(6):2371–2378. that these lines could be used in edamame Pennington. 1971. Stage of development de- Rao, M.S.S., A.S. Bhagsari, and A.I. Mohamed. breeding and/or commercial production. Fur- scriptions for soybeans, Glycine max (L.). 2002. Fresh green seed yield and seed nutri- ther evaluation is needed to confirm their Merrill. Crop Sci. 11:929–931. tional traits of vegetable soybean genotypes. superiority and release as cultivars and/or Holland, J.B. 2006. Estimating genotypic corre- Crop Sci. 42(6):1950–1958. lations and their standard errors using multi- Robertson, A. 1959. The sampling variance of the germplasm lines. variate restricted maximum likelihood genetic correlation coefficient. Biometrics estimation with SAS Proc MIXED. Crop Sci. 5:469–485. Literature Cited 46:642–654. Sharma, J.R. 1988. Statistical and biometrical Bernick, K. 2009. Edamame Takes Root in U.S. Kelley, K.M. and E.S. Sanchez. 2005. Accessing techniques in plant breeding. New Age In- Corn & Soybean Digest. 24 Feb. 2018. . 40:1347–1353. Shurtleff, W. and A. Aoyagi. 2009. History of Binder, K. 2010. Edible soybean rises in popularity Liu, K.S. 1999. Soybeans: Chemistry, technology, edamame, green vegetable soybeans, and with U.S. consumers & producers. Farm World. and utilization. Aspen Publishers, Inc., Gai- vegetable-type soybeans: Bibliography and 24 Apr. 2018. . McBryde, J. 2012. Snacking on Soy: Arkansas Lafayette, CA. Brar, G.S. and T.E. Carter, Jr. 1993. Soybean, Grows Edamame Commercially. Farm Flavor. Soyfoods Association of North America. 2011. p. 427–464. In: G. Kalloo and B.O. Bergh (eds.). 26 Feb. 2015. . history-of-soy-products>. Carson, L.C., J.H. Freeman, K. Zhou, G. Welbaum, Mebrahtu, T., T.E. Devine, P. Donald, and T.S. Williams, M.M., II. 2015. Phenomorphological and M. Reiter. 2011. Cultivar evaluation and Abney. 2005. Registration of ‘Asmara’ vege- characterization of vegetable soybean germ- lipid and protein contents of Virginia-grown table soybean. Crop Sci. 45:408–409. plasm lines for commercial production. Crop edamame. HortTechnology 21:131–135. Mebrahtu, T. and A. Mohamed. 2006. Genetic Sci. 55(3):1274–1279. CBS News. 2013. Edamame: Is the future of variation for green pod yield and quality among Zhang, L. and S. Kyei-Boahen. 2007. Growth and American soy farmers’ profits in edible soy vegetable soybean genotypes. J. Crop Improv. yield of vegetable soybean (Edamame) in beans for people? 24 Apr. 2018.

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