Characterization and Heterosis of High Quality Resistant Glutinous Rice Germplasms
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摘要:
目的 为有效改良传统糯稻品种的产量、品质及抗性等综合性状,满足当前专用型糯稻市场的发展需求。 方法 采用不完全双列杂交(NC II)方法,构建亲本及20个杂种一代的遗传群体,评价糯稻亲本及杂交组合产量和品质性状的育种利用价值,鉴定出高产、抗病、低直链淀粉含量、高碱消值的糯稻新种质。 结果 7个优良品系的实割产量变幅7744.35~9216.45 kg·hm−2,其中X-104实割产量最高,达9216.45 kg·hm−2,X-27实割产量达8960.40 kg·hm−2,明显优于3个对照;两个株系植株矮壮,且分别携带了Pi-1、Pi-kh和Pi-1、Pi-9、Pi-kh的稻瘟病抗性基因,表现为中抗稻瘟病;直链淀粉含量(amylose content, AC)分别为1.71%和1.68%;碱消值(alkali spreading value,ASV)达6.75级和6.13级,育种价值高。在杂种优势利用上,X-27除了株高性状一般配合力为正效应外,该株系有利提高杂种后代的产量与稻米品质。X-57降低后代的株高和AC,提高后代产量和ASV。X-104虽产量高,品质好,但不利于杂种优势利用。闽糯2S/X-41、闽糯2S/X-107杂交一代在产量、株高、AC及碱消值等性状上具有明显的超亲优势和竞争优势,在杂交糯稻上应用价值高。 结论 杂交糯稻的产量与品质均为复杂性状,其中有效穗数、穗长、千粒重、长宽比、单株产量、AC、ASV主要受糯稻恢复系的控制。各性状的广义遗传率(h2B)变幅14.72%~53.99%,狭义遗传率(h2N)变幅10.25%~43.07%,遗传率较低,易受环境影响。育种上需大量配组,方可培育出不同用途的专用型糯稻杂交稻。 Abstract:Objective In order to improve the yield, quality and resistance of traditional glutinous rice varieties, and meet the development demand of special glutinous rice market. Method Glutinous rice germplasms of high yield, disease resistance, low amylose content (AC), and high alkali spreading value (ASV) were evaluated and identified. Using the incomplete diallel crossing(NC II) method, the selected parental lines and 20 hybrids were pooled to construct a matrix for breeding evaluation. Result The actual yields of 7 choice germplasms ranged from 7744.35 to 9216.45 kg·hm−2, among which X-104 had the highest real cut yield of 9216.45 kg·hm−2 and X-27 had the highest real cut yield of 8960.40 kg·hm−2, which were significantly higher than those of the 3 reference varieties. The abovementioned two short and stout cultivars respectively carried Pi-1 and Pi-kh and Pi-1, Pi-9, and Pi-kh genes were moderate resistance to rice blast. They had AC of 1.71% and 1.68% as well as ASV of 6.75 and 6.13, respectively, indicating low gelatinization temperature. For heterosis in breeding, X-27 exhibited a combining ability of higher on grain yield and quality in addition to taller on plant height; X-57 of greater on yield and ASV but lower on plant height and AC; and X-104, despite the high grain yield and quality, did not offer meaningful advantages. Among the hybrids, Minnuo 2S/X-41 and Minnuo 2S/X-107 were significantly superior to their parents in terms of grain yield, plant height, AC, and ASV. Conclusion The determination of grain yield and quality of glutinous rice involves numerous factors such as effective panicle number, panicle length, 1000-grain weight, length-to-width ratio, yield per plant, AC, and ASV. The broad heritability (h2B) of such traits of the parents and hybrids in this study ranged between 14.72%–53.99%, and the narrow heritability (h2N) between 10.25%–43.07%. The relatively low indexes indicated instability and being prone to environmental effects in gene transfer from generation to generation. Consequently, breeding glutinous rice targeted for a specific use would require a complicated process of selection and hybridization. -
Key words:
- glutinous rice /
- amylose content /
- blast resistance /
- characterization /
- heterosis
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表 1 与3个抗病基因紧密连锁的引物序列
Table 1. Sequences of primers closely linked to 3 resistance genes
标记名称
Tag name引物序列(5′-3')
Primer sequence连锁的抗病基因
Linked resistance genes退火温度Tm/℃ RM224 F:ATCGATCGATCTTCACGAGG Pi-1 55 R:TGCTATAAAAGGCATTCGGG FP1+FP2+RP FP1:TCCCTGATTCCAACCTGCAGCAAGAG Pi-9 60 FP2:CCAACCTGCACCAGCCT RP:CCATGCTGATACTTGGGGCA RM206 F:CCCATGCGTTTAACTATTCT Pi-kh 55 R:CGTTCCATCGATCCGTATGG 表 2 7个糯稻株系稻瘟病抗性及品质性状
Table 2. Blast resistance and quality traits of 7 glutinous rice lines
株系
Strain携带抗病基因
Disease
resistance genes苗期叶瘟
Leaf plague at
seedling stage抗感类型
Type of
resistance碱消值
ASVX-27 Pi-1/Pi-kh 3 中抗 MR 6.75 X-41 Pi-9 3 中抗 MR 5.75 X-44 Pi-1 3 中抗 MR 6.38 X-57 Pi-1 Pi-9 3 中抗 MR 5.88 X-83 Pi-1 3 中抗 MR 5.38 X-104 Pi-1/Pi-9/Pi-kh 3 中抗 MR 6.13 X-107 Pi-1/Pi-9/Pi-kh 3 中抗 MR 5.88 珍珠糯(CK1) / 7 感 S 5.88 金糯6号(CK2) / 3 中抗 MR 5.25 金贵丝苗(CK3) / 4 中感 MS 3.88 表 3 7个优异品系的产量分析
Table 3. Yields and ACs of 7 premium rice lines
株系
Strain产量 Yield/(kg·hm−2) 直链淀粉含量AC/% 理论产量
Theoretical yield实割产量
Solid cut yield均值±标准差
Mean ± standard deviation变异系数
Coefficient of variation/%X-27 9542.85±1304.85abAB 8960.40±389.25aAB 1.71±0.23bcB 4.95 X-41 8368.65±994.80bcABC 8085.75±195.45bcCD 1.75±0.02bcB 1.35 X-44 8275.65±2164.50bcBC 7744.35±389.40cdCD 1.99±0.22bB 10.89 X-57 8484.15±1111.80bcABC 8107.05±266.40bcCD 1.82±0.07bcB 3.92 X-83 8285.55±1187.85bcBC 7808.40±230.85cdCD 1.81±0.07bcB 4.11 X-104 10164.45±2112.90aA 9216.45±64.05aA 1.68±0.08cB 4.72 X-107 8809.20±847.35abcABC 8320.50±230.70bBC 1.87±0.12bcB 6.64 珍珠糯(CK1) 8481.45±1131.75bcABC 8192.40±64.05bcCD 1.83±0.12bcB 6.65 金糯6号(CK2) 7689.60±1935.45cC 7531.05±37.05dD 1.92±0.12bcB 6.28 金贵丝苗(CK3) 8705.70±1241.40bcABC 8213.70±551.85bcC 16.95±0.50aA 2.98 大、小写字母分别表示0.01和0.05水平差异显著。
Data with upper- and lower-case letters indicate significant differences at P<0.01 and P<0.05, respectively.表 4 7个优异株系及对照品种的部分农艺性状
Table 4. Agronomic traits of 7 premium rice lines and reference varieties
株系
Strain株高
Plant height/
cm主穗长
Main panicle
length/cm主穗重
Main panicle
weight/g每穗总粒数
Total number of
grains per panicle粒长
Grain length/
mm长宽比
Length-width
ratio结实率
Setting
percentage/%千粒重
Thousand seed
weight/g单株产量
Yield per
plant/gX-27 101.3±1.9f 20.71±0.19e 5.33±0.33c 287.63±42.28ab 9.32±0.28de 3.96±0.27bcd 89.77±1.94bcd 21.81±1.05c 34.46±3.52bcd X-41 106.1±1.1def 22.95±0.64cd 5.28±0.34c 256.50±22.70bc 9.29±0.29e 3.98±0.20abcd 89.26±1.09bcd 21.88±0.92c 37.31±2.80b X-44 102.3±3.9ef 21.65±0.56de 4.72±0.64d 270.13±49.12b 9.38±0.24cde 4.06±0.28abc 90.64±1.20ab 22.20±0.74c 34.45±6.07bcd X-57 101.3±3.7f 21.16±0.44e 5.83±0.35ab 278.88±31.55ab 9.39±0.19cde 4.05±0.28abc 88.50±2.56bcd 21.76±0.97c 35.43±3.58bc X-83 103.4±2.3def 21.39±0.73de 5.25±0.60c 268.38±37.20b 9.60±0.20bc 4.14±0.32ab 87.51±2.22d 22.68±1.39c 33.51±3.38cd X-104 107.8±1.8cd 23.29±1.12c 6.07±0.77a 306.88±42.16a 9.54±0.32bcd 4.16±0.19ab 87.70±0.98d 22.53±1.21c 40.67±4.37a X-107 106.7±3.0 de 21.83±0.72cde 5.29±0.31c 271.50±29.86b 9.66±0.30b 4.27±0.40a 88.30±2.32cd 22.59±1.45c 36.48±1.94bc 珍珠糯
(CK1)128.6±4.4a 28.42±0.94a 5.92±0.53ab 211.01±25.60d 9.53±0.20bcd 3.82±0.21cd 89.27±2.89bcd 30.05±2.25a 32.04±1.64de 金糯6号
(CK2)118.2±3.8b 25.01±0.94b 5.44±0.54bc 226.88±17.08cd 9.54±0.06bcd 3.72±0.23d 92.61±2.96a 25.07±1.27b 28.79±3.98e 金贵丝苗
(CK3)112.5±3.5c 24.97±1.86b 4.63±0.64d 214.38±35.64d 10.21±0.26a 4.09±0.54abc 90.37±4.08abc 24.74±1.64b 36.79±2.31bc 同一性状不同小写字母表示差异达显著水平(P<0.05)。
Data with different lowercase letters on same trait indicate differences at P<0.05.表 5 亲本及其杂种后代主要产量及品质性状配合力方差
Table 5. Variance of combining ability on yield and quality traits of parents and their hybrid offspring
变异来源
Source of variation组合
Combination不育系一般
配合力方差 A恢复系一般
配合力方差 R组合特殊
配合力方差 A×R误差
Error株高
Plant height51.77** 70.73** 150.00** 14.28 13.15 有效穗数
Effective tiller number2.85** 5.96 2.59 2.17** 0.65 穗长
Panicle length5.92** 13.25 5.30 4.29 3.40 结实率
Setting percentage67.67** 140.46* 124.65* 30.48** 13.16 千粒重
Thousand seed weight3.50** 11.92** 3.82 1.29* 0.59 长宽比
Length-width ratio0.12** 0.37** 0.07 0.07 0.05 单株产量
Yeld per plant102.37** 327.69* 21.36 73.04** 22.83 直链淀粉含量AC 0.04** 0.10** 0.06* 0.02** 0.01 碱消值
ASV2.84** 3.68 8.09** 1.25* 0.67 *代表P<0.05的显著水平,**代表P<0.01的显著水平。
*: significant at P<0.05; **: significant at P<0.01.表 6 亲本主要产量和品质性状的GCA效应值
Table 6. GCA effect on yield and quality traits of parents
亲本
Parent株高
Plant height有效穗数
Effective tiller number穗长
Ear length结实率
Setting percentage千粒重
Thousand seed weight长宽比
Length-width ratio单株产量
Yield per plant直链淀粉
含量AC碱消值
ASVX-27 4.30 3.21 0.60 6.04 0.31 −1.63 4.64 −3.92 7.02 X-41 0.59 −7.78 −0.71 −7.12 5.33 −1.51 −4.68 3.22 1.75 X-57 −2.20 4.06 −3.49 0.81 −2.28 0.89 3.19 −9.46 1.65 X-104 2.02 −5.25 4.78 2.77 −0.87 −1.76 −2.03 6.46 −24.56 X-107 −4.71 5.75 −1.18 −2.49 −2.50 4.02 −1.12 3.69 15.79 糯326S −1.60 11.68 2.47 3.59 −4.92 1.26 19.84 2.26 −1.75 闽农糯6A −0.07 −7.28 3.75 −2.48 −0.44 −7.66 −9.18 −2.47 9.47 明糯208A 3.71 −3.89 −0.24 −5.51 7.00 4.07 −7.37 5.27 −27.02 闽糯2S −2.04 −0.51 −5.98 4.41 −1.65 2.33 −3.29 −5.06 19.30 表 7 20个组合主要产量和品质性状的SCA效应值
Table 7. SCA effect on yield traits, AC, and ASV in 20 combinations
组合
Combination株高
Plant
height有效穗数
Effective
tiller
number穗长
Ear
length结实率
Setting
percentage千粒重
Thousand
seed
weight长宽比
Length-width
ratio单株产量
Yield per
plant直链淀粉
含量AC碱消值
ASV糯326S/X-27 4.31 11.68 3.30 9.45 −9.21 −2.78 34.04 −0.85 12.28 闽农糯6A/X-27 5.58 −8.63 4.84 2.99 1.56 −7.18 −6.69 5.38 −1.75 明糯208A/X-27 7.16 15.06 1.52 −0.39 7.81 9.82 −4.80 0.65 −1.75 闽糯2S/X-27 6.94 −5.25 −7.26 12.11 1.09 −6.40 −4.01 −5.18 19.30 糯326S/X-41 0.58 1.52 6.02 0.76 3.43 4.99 0.12 4.22 −15.79 闽农糯6A/X-41 0.89 −22.17 4.72 −6.93 3.97 −12.96 −23.58 −0.63 12.28 明糯208A/X-41 1.26 −12.01 0.69 −22.79 10.53 −3.73 −0.95 −5.36 −29.82 闽糯2S/X-41 −3.94 1.52 −14.26 0.46 3.37 5.68 5.68 1.78 40.35 糯326S/X-57 −5.00 21.83 1.76 4.11 −1.14 4.73 30.17 1.49 19.30 闽农糯6A/X-57 −4.71 −5.25 −3.34 −5.40 −5.44 −8.04 −4.14 −0.72 19.30 明糯208A/X-57 3.68 1.52 −2.28 4.04 3.31 2.14 −0.03 −0.01 −29.82 闽糯2S/X-57 −6.22 −1.86 −10.10 0.49 −5.84 4.73 −13.23 −0.76 −8.77 糯326S/X-104 −2.41 1.52 −1.21 2.52 −6.10 −4.59 10.09 0.23 −43.86 闽农糯6A/X-104 2.69 −8.63 7.09 4.23 2.83 −4.68 −6.57 −11.57 −15.79 明糯208A/X-104 2.72 −25.55 1.99 −3.09 5.78 0.16 −29.08 6.05 −43.86 闽糯2S/X-104 1.49 11.68 11.24 7.42 −5.97 2.05 17.46 5.30 5.26 糯326S/X-107 −7.40 21.83 2.47 1.10 −11.57 3.95 24.76 −5.08 19.30 闽农糯6A/X-107 −5.37 8.29 5.43 −7.30 −5.11 −5.45 −4.90 7.54 33.33 明糯208A/X-107 2.26 1.52 −3.11 −5.33 7.59 11.97 −2.00 −1.32 −29.82 闽糯2S/X-107 −4.50 −8.63 −9.51 1.56 −0.89 5.59 −22.35 −1.14 40.35 表 8 杂种一代的产量和品质性状的超亲优势和竞争优势
Table 8. Yield, AC, and ASV of F1 hybrids that surpassed parents
组合
Combination单株重 Weight per plant 直链淀粉含量AC 碱消值ASV 超亲优势
Heterobeltiosis竞争优势CK1
Competitive
advantage
CK1竞争优势CK2
Competitive
advantage
CK2超亲优势
Heterobeltiosis竞争优势CK1
Competitive
advantage
CK1竞争优势CK2
Competitive
advantage
CK2超亲优势
Heterobeltiosis竞争优势CK1
Competitive
advantage
CK1竞争优势CK2
Competitive
advantage
CK2糯326S/X-27 38.40 55.15 74.12 1.17 −5.46 −9.90 −21.04 −16.46 −9.35 闽农糯6A/X-27 15.54 24.51 45.36 2.92 −3.83 −8.33 −30.81 −26.80 −20.58 明糯208A/X-27 17.89 32.16 48.32 5.85 −1.09 −5.73 −30.81 −26.80 −20.58 闽糯2S/X-27 18.88 33.27 49.57 −11.11 −16.94 −20.83 −16.00 −11.13 −3.57 糯326S/X-41 32.52 39.00 55.99 11.43 6.56 1.56 −30.43 −37.30 −31.97 闽农糯6A/X-41 1.15 6.10 19.07 1.71 −2.73 −7.29 −7.30 −16.46 −9.35 明糯208A/X-41 31.10 37.51 54.32 4.57 0.00 −4.69 −42.09 −47.81 −43.37 闽糯2S/X-41 39.86 46.70 64.64 1.14 −3.28 −7.81 16.00 4.55 13.44 糯326S/X-57 18.57 49.77 68.08 −8.24 9.29 −13.02 −3.57 −11.13 −3.57 闽农糯6A/X-57 5.37 33.09 49.36 −14.84 −15.30 −19.27 −3.57 −11.13 −3.57 明糯208A/X-57 9.88 38.78 55.75 −6.59 −7.10 −11.46 −43.37 −47.81 −43.37 闽糯2S/X-57 −4.63 20.46 35.19 −17.58 −18.03 −21.88 −26.36 −32.13 −26.36 糯326S/X-104 29.71 52.83 71.52 14.88 5.46 0.52 −56.44 −58.15 −54.59 闽农糯6A/X-104 10.09 29.71 45.57 −2.38 −10.38 −14.58 −34.75 −37.30 −31.97 明糯208A/X-104 −16.44 −1.55 10.49 24.40 14.21 8.85 −56.44 −58.15 −54.59 闽糯2S/X-104 38.40 63.08 83.01 12.50 3.28 −1.56 −18.43 −21.63 −14.97 糯326S/X-107 17.39 42.25 59.64 −4.28 −2.19 −6.77 −3.57 −11.13 −3.57 闽农糯6A/X-107 8.97 32.03 48.18 3.21 5.46 0.52 7.65 −0.78 7.65 明糯208A/X-107 12.29 36.06 52.69 2.14 4.37 −0.52 −43.37 −47.81 −43.37 闽糯2S/X-107 −11.03 7.80 20.98 −7.49 −5.46 −9.90 13.44 4.55 13.44 变化幅度 −16.44~39.86 −1.55~63.08 10.49~83.01 −17.58~24.40 −18.03~14.21 −21.88~8.85 −56.44~16.00 −58.15~4.55 −54.59~13.44 CK1为珍珠糯、CK2金糯6号。
CK1: Zhenzhunuo; CK2: Jinnuo 6.表 9 主要农艺性状及品质性状的遗传方差分量及遗传率
Table 9. Genetic variance components and heritability of major agronomic traits and AC
性状
Character基因型方差
Genotypic variance方差贡献率
Variance contribution rate/%遗传率
Heritability/%G1 G2 G12 误差
ErrorVg Vg1 Vg2 Vs h2B h2N 株高
Plant height6.786 2.258 0.226 13.154 97.6 73.2 24.4 2.4 41.34 40.33 有效穗数
Effective tiller number0.021 0.152 0.303 0.650 36.3 4.4 31.9 63.7 42.28 15.34 穗长
Panicle length0.051 0.358 0.178 3.401 69.7 8.6 61.1 30.3 14.72 10.25 结实率
Setting percentage4.709 4.399 3.463 13.164 72.5 37.5 35.0 27.6 48.85 35.39 千粒重
Thousand seed weight0.126 0.425 0.140 0.589 79.8 18.3 61.5 20.2 53.99 43.07 长宽比
Length-width ratio0.009 0.012 0.004 0.050 76.4 36.4 48.6 23.6 23.99 18.33 单株产量
Yeld per plant2.253 10.186 10.043 22.828 50.4 10.0 45.3 49.7 46.98 23.66 AC 0.002 0.004 0.002 0.007 71.1 21.5 49.4 28.9 51.95 36.95 ASV 0.121 0.273 0.116 0.671 77.4 23.7 53.5 22.6 43.20 33.43 G1表示糯稻不育系GCA基因型方差,G2表示糯稻恢复系GCA基因型方差,G12表示糯稻不育系与糯稻恢复系互作(SCA)基因型方差,Vg表示GCA基因型方差占总方差比,Vg1表示糯稻不育系GCA基因型方差占总方差比,Vg2表示糯稻恢复系GCA基因型方差占总方差比,Vs表示糯稻不育系和糯稻恢复系互作(SCA)基因型方差占总方差比。h2B表示广义遗传率,h2N表示狭义遗传率。
G1: Variance of GCA genotypes of glutinous sterile lines; G2: variance of GCA genotypes of glutinous recovery lines; G12: variance of SCA genotypes of glutinous sterile lines and glutinous recovery lines; Vg: variance of GCA genotypes to total variance; Vg1: variance of GCA genotypes of glutinous sterile lines to total variance; Vg2: variance of GCA genotypes of glutinous recovery lines to total variance; Vs: variance of SCA genotypes of glutinous sterile lines to total variance; h2B: broad heritability; h2N: narrow heritability. -
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