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作物学报

作物学报 ›› 2024, Vol. 50 ›› Issue (5): 1271-1286.doi: 10.3724/SP.J.1006.2024.34134

• 耕作栽培·生理生化 • 上一篇    下一篇

外源物质浸种对迟播油菜越冬期抗寒性及产量的影响

王先领(), 姜岳, 雷贻忠, 肖胜男, 厍惠洁, 段圣省, 黄铭, 蒯婕, 汪波, 王晶, 赵杰, 徐正华*(), 周广生   

  1. 华中农业大学植物科学技术学院 / 农业农村部长江中游作物生理生态与耕作重点实验室, 湖北武汉 430070
  • 收稿日期:2023-08-05 接受日期:2024-01-12 出版日期:2024-05-12 网络出版日期:2024-02-09
  • 通讯作者: 徐正华, E-mail: xzh@mail.hzau.edu.cn
  • 作者简介:E-mail: xlwang@webmail.hzau.edu.cn
  • 基金资助:
    湖北省重点研发计划项目(2023BBB028);财政部和农业农村部国家现代农业产业技术体系建设专项(CARS-12)

Effects of seed soaking with exogenous substances on late-seeded rapeseed cold resistance of during overwintering period and yield

WANG Xian-Ling(), JIANG Yue, LEI Yi-Zhong, XIAO Sheng-Nan, SHE Hui-Jie, DUAN Sheng-Xing, HUANG Ming, KUAI Jie, WANG Bo, WANG Jing, ZHAO Jie, XU Zheng-Hua*(), ZHOU Guang-Sheng   

  1. College of Plant Science and Technology, Huazhong Agricultural University / Key Laboratory of Crop Ecophysiology and Farming System for the Middle Reaches of the Yangtze River, Ministry of Agriculture and Rural Affairs, Wuhan 430070, Hubei, China
  • Received:2023-08-05 Accepted:2024-01-12 Published:2024-05-12 Published online:2024-02-09
  • Contact: E-mail: xzh@mail.hzau.edu.cn
  • Supported by:
    Key Research and Development Plan of Hubei Province(2023BBB028);China Agriculture Research System of MOF and MARA(CARS-12)

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摘要：

确保迟播油菜产量是压减长江流域双季稻区冬闲田的关键, 提高越冬期抗寒性、促进冬前干物质积累是增加迟播油菜产量的有效途径。播前外源物质浸种是提高越冬期抗寒性, 促进冬前快速生长的有效措施。试验选用早熟品种华油杂137, 设置清水(CK), 0.01 mmol L-1、0.05 mmol L-1、0.10 mmol L-1的甜菜碱(T1-1、T1-2、T1-3), 0.1 mmol L-1、0.5 mmol L-1、1.0 mmol L-1的脯氨酸(T2-1、T2-2、T2-3), 0.03%、0.15%、0.30%的过氧化氢(T3-1、T3-2、T3-3), 0.001 mmol L-1、0.01 mmol L-1、0.05 mmol L-1的苹果酸(T4-1、T4-2、T4-3), 25 mg L-1、100 mg L-1、300 mg L-1的氧化纳米锌(T5-1、T5-2、T5-3), 0.5 mmol L-1、1.0 mmol L-1的多胺(T6-1、T6-2)浸种处理后, 于2021—2023年开展田间试验, 研究不同外源物质及浓度浸种对迟播油菜越冬期抗寒性及产量的影响。结果表明, 不同外源物质和不同浓度浸种处理影响迟播油菜出苗率, 部分处理如T3-3、T4-2、T5-3与CK相比显著提高, 提高幅度分别达到19.2%、15.3%、17.3%。部分外源物质浸种也显著提高了迟播油菜抗寒性, 其机制一方面是提高抗氧化系统中过氧化物酶、过氧化氢酶活性和谷胱甘肽含量, 减少活性氧、过氧化氢及丙二醛含量, 另一方面是增加可溶性糖、脯氨酸、苹果酸、多胺、甘露醇等渗透调节物质含量, 提高膜联冷应答蛋白激酶含量从而提高抗寒性。叶片抗寒性的增加有利于越冬期叶片生物量的积累, 有效分枝数增加, 单株角果数增加, 从而促进产量的提高。另外, 对筛选出的9个正向指标和3个负向抗寒性指标进行综合评价, 结果表明, 2年平均产量排序和抗寒性综合评价值排序结果基本一致, 产量较高、抗寒性较强的浸种处理为T5-3、T5-2、T2-3、T1-2。该研究可为长江流域迟播油菜冬前壮苗培育、提高越冬期抗寒性提供理论与技术支撑, 为迟播油菜抗逆稳产、促进冬闲田开发利用提供依据。

关键词: 油菜, 迟播, 抗寒性, 外源物质, 浸种

Abstract:

In double-cropping rice areas in the Yangtze River Basin (YRB), ensuring the yield of late-seeded rapeseed is essential to reduce the winter fallow fields. Additionally, promoting the accumulation of dry matter before winter and improving cold resistance during the overwintering period are effective ways to increase the yield of late-seeded rapeseed. Soaking seeds with exogenous substances is an effective measure to improve the cold resistance during the overwintering period and promote the rapid growth before winter. In this experiment, the early maturing rapeseed variety Huayouza 137 was selected, while soaking-seed treatments of water (CK); 0.01 mmol L-1, 0.05 mmol L-1, 0.10 mmol L-1 betaine (T1-1, T1-2, T1-3); 0.1 mmol L-1, 0.5 mmol L-1, 1.0 mmol L-1 proline (T2-1, T2-2, T2-3); 0.03%, 0.15%, 0.30% hydrogen peroxide (T3-1, T3-2, T3-3); 0.001 mmol L-1, 0.01 mmol L-1, 0.05 mmol L-1 malic acid (T4-1, T4-2, T4-3); 25 mg L-1, 100 mg L-1, 300 mg L-1 nano zinc oxide (T5-1, T5-2, T5-3); 0.5 mmol L-1, 1.0 mmol L-1 polyamines (T6-1, T6-2) were conducted between 2021 and 2023 rapeseed growing season. We studied the effects of soaking seeds with different exogenous substances and their levels on the cold resistance in winter and yield of late-seeded rapeseed. The results showed that the different exogenous substances and their levels of soaking seeds affected the germination rate of late-seeded rapeseed, and part of them, such as T3-3, T4-2, and T5-3, significantly increased by 19.2%, 15.3%, and 17.3% versus CK. Soaking seeds with some exogenous substances significantly improved the cold resistance of late-seeded rapeseed in winter. On the one hand, the activities of peroxidase, catalase, and the content of glutathione were improved, and the contents of hydrogen peroxide, active oxygen and MDA reduced; on the other hand, the contents of soluble sugar, proline, malic acid, polyamine, and mannitol were increased; at the same time, the content of membrane cold response protein kinase was increased. The increase of leaf cold resistance was beneficial to the accumulation of leaf biomass and the increase of effective branches and pod number per plant, thus promoting yield. In addition, 9 positive and 3 negative cold resistance indicators were comprehensively evaluated. The results showed that the two-year average yield ranking and the comprehensive evaluation value ranking of cold resistance were basically the same over the two years. While the soaking-seed treatments with the best yield and cold resistance were T5-3, T5-2, T2-3, and T1-2. These results of this study provide theoretical and technical support for the cultivation of late-seeded rapeseed seedlings before winter and the improvement of cold resistance in the YRB, and provide a basis for the stress resistance and stable production of late-seeded rapeseed and the development and utilization of winter fallow fields.

Key words: rapeseed, late-seeded, cold resistance, exogenous substance, soaking seeds

表1

用于迟播油菜浸种的不同外源物质及浓度"

浓度
Concentration		 对照
Contrast
(CK)		 甜菜碱
Betaine
(T1)		 脯氨酸
Proline
(T2)		 过氧化氢
Hydrogen peroxide (T3)		 苹果酸
Malic acid
(T4)		 氧化纳米锌
Nano zinc oxide
(T5)		 多胺
Polyamine
(T6)
浓度-1
Concentration-1		 — 0.01 mmol L-1 0.1 mmol L-1 0.03% 0.001 mmol L-1 25 mg L-1 0.5 mmol L-1
浓度-2
Concentration-2		 — 0.05 mmol L-1 0.5 mmol L-1 0.15% 0.01 mmol L-1 100 mg L-1 1.0 mmol L-1
浓度-3
Concentration-3		 — 0.10 mmol L-1 1.0 mmol L-1 0.30% 0.05 mmol L-1 300 mg L-1 —

图1

2021-2022生长季(a)和2022-2023生长季(b)的主要气象因子 Tmin和Tmax分别代表日最低温和日最高温。"

图2

不同外源物质浸种对迟播油菜2021-2022 (a)和2022-2023 (b)生长季出苗率的影响 不同小写字母表示不同外源物质浸种处理间差异达显著水平(P < 0.05)。处理同表1。"

表2

不同外源物质浸种对迟播油菜产量及产量构成的影响"

年份
Year		 外源物质
Exogenous
substance		 浓度
Concentration		 单株角果数
Pod number per plant		 每角粒数
Seed number per pod		 千粒重1000-seed weight (g) 单株产量
Yield
per plant (g)		 成株率Survival rate (%) 产量
Yield
(kg hm-2)
2021-
2022		 对照CK CK 128.7 ef 17.40 bc 3.55 ab 7.95 bc 45.27 d 2347.1 gh
甜菜碱
Betaine (T1)		 T1-1 125.1 ef 17.84 a 3.40 abc 7.61 bc 47.56 bcd 2112.9 i
T1-2 145.8 cde 17.31 c 3.60 a 9.09 ab 46.95 bcd 2558.7 abc
T1-3 146.3 cde 17.68 ab 3.44 abc 8.89 abc 47.33 bcd 2549.6 abc
脯氨酸
Proline (T2)		 T2-1 139.8 cdef 17.66 ab 3.51 abc 8.66 abc 47.18 bcd 2563.1 abc
T2-2 178.7 a 17.48 abc 3.37 bc 10.53 a 49.68 abc 2571.5 abc
T2-3 163.4 abc 17.42 bc 3.43 abc 9.75 ab 49.81 abc 2591.8 abc
过氧化氢
Hydrogen
peroxide (T3)		 T3-1 114.5 f 17.34 bc 3.45 abc 6.87 c 49.82 abc 1771.8 j
T3-2 126.7 ef 17.69 ab 3.52 abc 7.89 bc 47.17 bcd 2321.5 h
T3-3 134.1 def 17.46 abc 3.34 c 7.83 bc 47.58 bcd 2430.2 efg
苹果酸
Malic acid (T4)		 T4-1 136.1 def 17.26 c 3.47 abc 8.13 bc 49.65 abc 2444.2 ef
T4-2 141.5 cdef 17.38 bc 3.48 abc 8.57 abc 49.92 abc 2451.6 def
T4-3 138.4 cdef 17.66 ab 3.51 abc 8.56 abc 44.65 d 2419.8 fg
氧化纳米锌
Nano zinc oxide (T5)		 T5-1 159.8 abcd 17.32 bc 3.48 abc 9.66 ab 45.74 cd 2536.1 bcd
T5-2 163.8 abc 17.53 abc 3.34 c 9.60 ab 52.43 a 2603.6 ab
T5-3 171.7 ab 17.45 abc 3.51 abc 10.52 a 50.00 ab 2635.1 a
多胺
Polyamine (T6)		 T6-1 148.8 bcde 17.39 bc 3.44 abc 8.90 abc 47.21 bcd 2451.3 def
T6-2 135.1 def 17.67 ab 3.44 abc 8.20 bc 47.15 bcd 2510.4 cde
2022-
2023		 对照CK CK 126.0 de 17.19 b 3.54 ab 7.66 cde 47.09 ab 2280.0 e
甜菜碱
Betaine (T1)		 T1-1 122.0 de 17.44 ab 3.37 bc 7.20 de 50.45 a 2050.5 f
T1-2 139.4 cd 17.73 a 3.58 a 8.85 abcd 48.46 ab 2508.4 a
T1-3 141.9 bcd 17.67 a 3.33 bc 8.37 bcde 45.45 ab 2449.9 abcd
脯氨酸
Proline (T2)		 T2-1 133.0 d 17.16 b 3.47 abc 7.90 cde 50.05 ab 2482.5 abc
T2-2 174.2 a 17.52 ab 3.45 abc 10.54 a 46.19 ab 2488.5 ab
T2-3 160.7 abc 17.62 ab 3.31 c 9.38 abc 48.96 ab 2532.2 a
2022-
2023		 过氧化氢
Hydrogen
peroxide (T3)		 T3-1 108.3 e 17.44 ab 3.50 ab 6.62 e 46.27 ab 1671.1 g
T3-2 123.7 de 17.64 ab 3.50 ab 7.63 cde 46.81 ab 2261.2 e
T3-3 129.4 de 17.74 a 3.30 c 7.56 cde 48.36 ab 2366.2 bcde
苹果酸
Malic acid (T4)		 T4-1 129.5 de 17.39 ab 3.53 ab 7.97 cde 47.89 ab 2355.1 de
T4-2 134.1 d 17.56 ab 3.53 ab 8.33 bcde 47.61 ab 2359.4 cde
T4-3 134.7 d 17.57 ab 3.48 abc 8.21 bcde 49.18 ab 2339.2 de
氧化纳米锌
Nano zinc oxide (T5)		 T5-1 157.2 abc 17.37 ab 3.45 abc 9.44 abc 50.37 ab 2453.1 abcd
T5-2 159.2 abc 17.48 ab 3.48 abc 9.70 abc 47.66 ab 2525.9 a
T5-3 164.1 ab 17.55 ab 3.52 ab 10.14 ab 47.83 ab 2575.0 a
多胺
Polyamine (T6)		 T6-1 146.8 bcd 17.44 ab 3.44 abc 8.81 abcd 48.40 ab 2358.4 cde
T6-2 129.6 de 17.36 ab 3.51 abc 7.89 cde 45.10 b 2454.8 abcd
方差分析 ANOVA
年份 Year * ns ns ns ns **
处理 Treatment ** ns ns ** ns **
年份×处理 Year×Treatment ns ns ns ns ns ns

表3

不同外源物质浸种对迟播油菜成熟期关健农艺性状的影响"

年份
Year		 外源物质
Exogenous
substance		 浓度
Concentration		 株高
Plant
height (cm)		 有效分枝数Effective branch number 根颈粗
Root-crown
diameter (mm)		 地上部干重
Shoot
biomass (g)		 抗折力
Bending
resistance (N)
2021-2022 对照CK CK 164.1 ab 4.6 bcd 10.8 bcd 23.6 abc 79.2 g
甜菜碱
Betaine (T1)		 T1-1 171.5 ab 4.5 bcd 11.4 abcd 22.6 bc 64.2 h
T1-2 170.8 ab 5.9 a 10.0 d 29.7 abc 96.4 cd
T1-3 168.6 ab 5.6 abc 11.9 abcd 29.7 abc 95.4 cd
2021-2022 脯氨酸
Proline (T2)		 T2-1 174.6 a 5.6 ab 12.1 abc 30.0 abc 96.4 cd
T2-2 164.2 ab 5.1 abcd 11.1 abcd 30.7 ab 91.8 de
T2-3 170.4 ab 5.4 abcd 12.7 ab 31.1 ab 115.6 a
过氧化氢
Hydrogen
peroxide (T3)		 T3-1 166.8 ab 4.8 abcd 11.4 abcd 20.7 c 82.4 efg
T3-2 172.3 ab 5.3 abcd 12.8 a 23.5 abc 91.6 de
T3-3 157.8 b 5.2 abcd 11.1 abcd 24.7 abc 102.2 bc
苹果酸
Malic acid (T4)		 T4-1 173.1 a 5.5 abc 12.5 ab 24.8 abc 81.4 fg
T4-2 166.9 ab 4.9 abcd 12.2 abc 26.3 abc 94.4 cd
T4-3 169.6 ab 4.7 bcd 10.5 cd 24.7 abc 92.6 cd
氧化纳米锌
Nano zinc oxide (T5)		 T5-1 170.5 ab 4.4 cd 11.1 abcd 29.1 abc 81.4 fg
T5-2 170.3 ab 4.2 d 10.5 cd 32.4 a 94.4 cd
T5-3 166.3 ab 4.3 d 11.2 abcd 32.5 a 92.6 cd
多胺
Polyamine (T6)		 T6-1 173.7 a 5.5 abc 12.2 abc 27.2 abc 107.0 ab
T6-2 170.8 ab 4.8 abcd 11.1 abcd 27.6 abc 89.4 def
2022-2023 对照CK CK 166.3 ab 4.8 bcd 10.4 abc 22.6 abc 69.2 g
甜菜碱
Betaine (T1)		 T1-1 164.6 ab 4.4 d 10.3 bc 22.2 abc 54.2 h
T1-2 172.0 ab 4.5 cd 11.0 abc 20.9 bc 86.4 bcd
T1-3 171.4 ab 6.0 a 9.4 c 29.0 ab 85.4 cd
脯氨酸
Proline (T2)		 T2-1 169.1 ab 5.7 abc 11.2 abc 28.7 ab 86.4 bcd
T2-2 175.0 a 5.7 ab 11.5 ab 28.6 ab 81.8 de
T2-3 164.7 ab 5.2 abcd 10.6 abc 30.8 a 95.6 ab
过氧化氢
Hydrogen
peroxide (T3)		 T3-1 171.0 ab 5.5 abcd 12.3 a 29.5 ab 72.4 efg
T3-2 166.9 ab 4.9 abcd 10.5 abc 19.4 c 81.6 de
T3-3 172.5 ab 5.4 abcd 12.1 ab 22.5 abc 92.2 abc
苹果酸
Malic acid (T4)		 T4-1 158.3 b 5.3 abcd 10.7 abc 23.9 abc 71.4 fg
T4-2 173.7 ab 5.6 abc 11.8 ab 23.7 abc 84.4 cd
T4-3 167.4 ab 4.9 abcd 11.6 ab 25.7 abc 82.6 cd
氧化纳米锌
Nano zinc oxide (T5)		 T5-1 170.2 ab 4.8 abcd 10.1 bc 23.7 abc 71.4 fg
T5-2 170.9 ab 4.5 cd 10.3 bc 28.0 abc 84.4 cd
T5-3 170.5 ab 4.3 d 10.2 bc 27.9 abc 82.6 cd
多胺
Polyamine (T6)		 T6-1 166.9 ab 4.3 d 10.6 abc 31.6 a 97.0 a
T6-2 174.4 a 5.6 abc 11.7 ab 25.7 abc 79.4 def
方差分析 ANOVA
年份 Year ns ns ** ns **
处理 Treatment ns ** ns ** **
年份×处理 Year×Treatment ** ns ** ns ns

表4

不同外源物质浸种对迟播油菜越冬期关键农艺性状及叶片生物量的影响"

年份
Year		 外源物质
Exogenous
substance		 浓度
Concentration		 株高
Plant
Height (cm)		 根长
Root
length (cm)		 根颈粗
Root-crown
diameter (mm)		 叶面积
Leaf
area (cm2)		 叶片生物量
Leaf
biomass (g)
2021-
2022		 对照CK CK 22.4 b 12.2 abc 2.80 cde 436.6 de 1.24 gh
甜菜碱
Betaine (T1)		 T1-1 21.8 bc 11.9 bc 2.71 de 423.6 e 1.13 h
T1-2 21.0 bc 13.4 abc 3.43 abcd 487.9 bcde 1.63 cd
T1-3 21.0 bc 12.8 abc 3.37 abcde 481.1 bcde 1.55 de
脯氨酸
Proline (T2)		 T2-1 20.1 bc 14.1 a 3.57 abc 498.3 bcd 1.71 bc
T2-2 20.8 bc 12.8 abc 3.58 abc 506.9 bc 1.73 bc
T2-3 21.6 bc 12.9 abc 3.63 abc 507.1 bc 1.82 ab
过氧化氢
Hydrogen
peroxide (T3)		 T3-1 21.4 bc 12.6 abc 2.57 e 326.0 f 0.77 i
T3-2 21.7 bc 13.3 abc 2.80 cde 426.2 e 1.24 gh
T3-3 22.0 bc 12.9 abc 2.99 bcde 448.7 cde 1.38 efg
苹果酸
Malic acid (T4)		 T4-1 23.9 a 12.6 abc 3.07 abcde 460.6 cde 1.41 ef
T4-2 20.2 bc 13.1 abc 3.14 abcde 470.7 cde 1.46 e
T4-3 21.4 bc 11.9 bc 2.91 bcde 446.2 cde 1.27 fgh
氧化纳米锌
Nano zinc oxide (T5)		 T5-1 19.6 c 12.7 abc 3.27 abcde 476.0 bcde 1.49 e
T5-2 20.6 bc 13.6 abc 3.69 ab 537.0 ab 1.89 a
T5-3 21.1 bc 14.0 ab 3.84 a 571.7 a 1.93 a
多胺
Polyamine (T6)		 T6-1 20.7 bc 12.0 abc 3.10 abcde 470.1 cde 1.43 e
T6-2 20.8 bc 11.4 c 3.23 abcde 473.3 bcde 1.48 e
2022-
2023		 对照CK CK 22.3 ab 12.4 bc 2.95 abc 435.7 cd 1.16 de
甜菜碱
Betaine (T1)		 T1-1 21.3 abc 12.3 bc 2.84 bc 413.6 d 1.02 e
T1-2 20.3 bcd 14.3 abc 3.49 abc 481.2 bcd 1.46 bc
T1-3 20.8 bcd 13.2 abc 3.50 abc 473.0 bcd 1.49 bc
脯氨酸
Proline (T2)		 T2-1 19.5 cd 14.9 a 3.68 abc 488.9 bc 1.63 ab
T2-2 20.1 bcd 13.5 abc 3.78 ab 501.6 bc 1.61 ab
T2-3 21.2 abcd 13.2 abc 3.73 ab 493.5 bc 1.72 a
过氧化氢
Hydrogen
peroxide (T3)		 T3-1 21.0 bcd 13.1 abc 2.73 c 318.5 e 0.69 f
T3-2 21.4 abc 13.5 abc 2.89 bc 416.8 d 1.16 de
T3-3 21.5 abc 13.3 abc 3.17 abc 439.9 cd 1.21 de
苹果酸
Malic acid (T4)		 T4-1 23.1 a 12.8 abc 3.16 abc 446.5 cd 1.29 cd
T4-2 20.0 bcd 13.6 abc 3.22 abc 454.9 bcd 1.34 cd
T4-3 20.9 bcd 12.5 bc 3.08 abc 438.0 cd 1.15 de
氧化纳米锌
Nano zinc oxide (T5)		 T5-1 18.9 d 13.2 abc 3.33 abc 471.3 bcd 1.38 cd
T5-2 20.3 bcd 14.0 abc 3.79 ab 521.2 ab 1.75 a
T5-3 20.9 bcd 14.4 ab 3.93 a 559.9 a 1.77 a
多胺
Polyamine (T6)		 T6-1 20.2 bcd 12.3 bc 3.23 abc 459.1 bcd 1.32 cd
T6-2 20.4 bcd 12.0 c 3.41 abc 464.8 bcd 1.32 cd
方差分析 ANOVA
年份 Year ** ** * ns **
处理 Treatment ** ** ** ** **
年份×处理 Year×Treatment ns ns ns ns ns

图3

产量与越冬期迟播油菜(a)根颈粗、(b)叶面积和(c)叶片生物量的相关性分析"

表5

不同外源物质浸种对迟播油菜越冬期叶片活性氧及抗氧化系统的影响"

外源物质
Exogenous
substance		 浓度
Concentration		 MDA
(nmol g-1)		 H2O2
(μmol g-1)		 O2∙-
(μg g-1)		 ·OH
(ng g-1)		 SOD
(U)		 POD
(U)		 CAT
(U)		 APX
(U)
对照CK CK 184.3 a 0.983 b 5.56 b 7.37 a 1730.1 g 2.64 def 589.3 d 0.86 e
甜菜碱
Betaine (T1)		 T1-1 190.2 a 1.014 b 6.29 a 7.85 a 1727.4 g 2.48 f 569.8 d 1.17 cd
T1-2 121.7 cde 0.682 cd 5.48 bc 5.08 cd 2463.3 cd 3.52 bc 968.2 b 1.10 d
T1-3 128.6 cd 0.741 c 4.95 bcde 5.24 c 2319.4 cde 3.41 c 937.1 b 0.94 e
脯氨酸
Proline (T2)		 T2-1 116.6 cde 0.614 d 4.74 cde 4.69 cde 2485.1 cd 3.72 abc 969.1 b 0.91 e
T2-2 115.8 cde 0.589 d 5.33 bc 4.69 cde 2585.5 c 3.80 ab 982.7 b 1.35 ab
T2-3 115.3 cde 0.589 d 4.49 def 4.67 cde 2938.6 b 3.83 ab 1002.2 b 1.43 a
过氧化氢
Hydrogen
peroxide (T3)		 T3-1 192.8 a 1.123 a 4.43 ef 7.96 a 1091.9 h 1.59 g 561.1 d 0.85 e
T3-2 185.4 a 1.002 b 3.94 f 7.40 a 1730.1 g 2.56 ef 589.3 d 0.98 e
T3-3 177.9 a 0.968 b 5.18 bcd 7.16 a 1855.0 fg 2.84 def 681.6 cd 0.97 e
苹果酸
Malic acid (T4)		 T4-1 157.3 b 0.937 b 3.97 f 6.37 b 1998.9 efg 2.93 de 740.8 c 1.26 bc
T4-2 155.3 b 0.918 b 6.17 a 6.29 b 2069.6 defg 2.95 de 772.9 c 1.12 cd
T4-3 183.3 a 0.969 b 4.60 def 7.25 a 1768.1 fg 2.69 def 613.5 d 0.95 e
氧化纳米锌
Nano zinc oxide (T5)		 T5-1 134.2 c 0.773 c 5.50 bc 5.27 c 2210.8 cdef 3.08 d 917.6 b 0.98 e
T5-2 108.1 de 0.570 d 6.45 a 4.30 de 3047.2 ab 4.00 a 1013.8 b 1.15 cd
T5-3 105.4 e 0.561 d 6.23 a 4.14 e 3313.4 a 4.11 a 1122.6 a 0.90 e
多胺
Polyamine (T6)		 T6-1 138.5 c 0.891 b 5.33 bc 6.19 b 2199.9 cdef 3.00 de 780.6 c 1.21 cd
T6-2 135.2 c 0.781 c 4.77 cde 5.44 c 2191.8 cdef 3.00 de 891.4 b 1.15 cd

表6

不同外源物质浸种对迟播油菜越冬期叶片非酶促清除系统的影响"

外源物质
Exogenous substance		 浓度
Concentration		 抗坏血酸
ASA (μmol g-1)		 类胡萝卜素
Carnosine (pmol g-1)		 谷胱甘肽
GSH (mg g-1)
对照CK CK 6.91 b 1.22 def 4.06 d
甜菜碱
Betaine (T1)		 T1-1 5.76 c 1.19 ef 3.46 e
T1-2 9.13 a 1.03 gh 4.88 bc
T1-3 7.14 b 0.96 h 4.84 bc
脯氨酸
Proline (T2)		 T2-1 7.66 b 1.17 f 5.07 ab
T2-2 5.57 c 1.53 ab 5.14 ab
T2-3 9.42 a 1.29 cde 5.23 ab
过氧化氢
Hydrogen peroxide (T3)		 T3-1 8.88 a 1.10 fg 3.28 e
T3-2 8.73 a 1.12 fg 3.47 e
T3-3 6.86 b 1.33 cd 4.34 cd
苹果酸
Malic acid (T4)		 T4-1 5.90 c 1.37 c 4.37 cd
T4-2 5.51 c 1.29 cde 4.44 cd
T4-3 7.22 b 1.51 ab 4.11 d
氧化纳米锌
Nano zinc oxide (T5)		 T5-1 7.31 b 1.35 cd 4.83 bc
T5-2 8.65 a 1.42 bc 5.48 a
T5-3 5.84 c 1.30 cde 5.56 a
多胺
Polyamine (T6)		 T6-1 6.86 b 1.38 c 4.79 bc
T6-2 9.77 a 1.56 a 4.81 bc

表7

不同外源物质浸种对迟播油菜越冬期叶片渗透调节物质的影响"

外源物质
Exogenous
substance		 浓度
Concentration		 可溶性糖
Soluble sugar
(mg g-1)		 脯氨酸
Proline
(ng g-1)		 苹果酸
Malate
(μg g-1)		 甜菜碱
Betaine
(pg g-1)		 多胺
Polyamine
(mg g-1)		 甘露醇
Mannitol
(ng g-1)
对照CK CK 4.28 k 268.5 b 3.71 f 398.6 cd 5.69 ef 126.7 e
甜菜碱
Betaine (T1)		 T1-1 4.11 l 211.7 c 3.40 f 475.9 ab 5.45 ef 125.0 e
T1-2 5.15 ef 319.3 a 5.03 bcd 365.0 cde 9.24 ab 181.8 bcd
T1-3 5.13 ef 308.8 a 4.82 cd 523.3 a 8.99 abc 177.6 bcd
脯氨酸
Proline (T2)		 T2-1 5.27 d 318.0 a 5.06 bcd 319.2 e 9.31 ab 184.4 bcd
T2-2 5.23 de 321.8 a 5.17 bc 338.5 de 9.71 a 186.4 bc
T2-3 5.39 c 318.7 a 5.42 bc 383.0 cde 9.32 ab 187.9 bc
过氧化氢
Hydrogen
peroxide (T3)		 T3-1 4.05 l 256.6 b 2.89 g 488.2 ab 5.01 f 70.4 f
T3-2 4.27 k 303.8 a 3.48 f 492.7 ab 5.52 ef 125.5 e
T3-3 4.42 j 264.7 b 3.78 f 483.7 ab 5.97 ef 159.2 cd
苹果酸
Malic acid (T4)		 T4-1 4.55 i 215.0 c 3.81 f 354.4 de 6.10 ef 161.4 cd
T4-2 4.72 h 253.0 b 4.06 ef 473.0 ab 7.91 d 162.9 cd
T4-3 4.41 j 267.7 b 3.67 f 325.8 e 5.79 ef 153.6 d
氧化纳米锌
Nano zinc oxide (T5)		 T5-1 5.04 f 300.0 a 4.77 cd 388.4 cde 8.12 cd 177.1 bcd
T5-2 5.60 b 330.9 a 5.59 ab 428.4 bc 10.02 a 201.8 ab
T5-3 6.31 a 322.8 a 6.00 a 369.1 cde 10.04 a 215.5 a
多胺
Polyamine (T6)		 T6-1 4.84 g 323.3 a 4.15 ef 513.9 a 6.61 e 165.8 cd
T6-2 5.04 f 264.7 b 4.45 de 512.7 a 8.45 bcd 168.7 cd

图4

不同外源物质浸种对迟播油菜越冬期叶片膜联冷应答蛋白激酶和Ca2+通道蛋白的影响 不同小写字母表示不同外源物质浸种处理间差异达显著水平(P < 0.05)。MRPK: 膜联冷应答蛋白激酶; VGCC: Ca2+通道蛋白。处理同表1。"

图5

抗寒性生理指标的主成分分析 Y: 产量; LA: 叶面积; LB: 叶片生物量; POD: 过氧化物酶; SOD: 超氧化物歧化酶; CAT: 过氧化氢酶; GSH: 谷胱甘肽; MRPK: 膜联冷应答蛋白激酶; Mal: 苹果酸; Pol: 多胺; Man: 甘露醇; ASA: 抗坏血酸; Car: 类胡萝卜素; O2?-: 超氧阴离子自由基; SS: 可溶性糖; Pro: 脯氨酸; Bet: 甜菜碱; VGCC: Ca2+通道蛋白; APX: 抗坏血酸过氧化物酶; H2O2: 过氧化氢; ?OH: 羟自由基; MDA: 丙二醛。"

表8

迟播油菜外源物质浸种抗寒性Topsis综合评价及平均产量排序结果"

外源物质
Exogenous
substance		 评价对象
Evaluation
object		 正理想解距离D+
Positive ideal solution distance D+		 负理想解距离D- Negative ideal solution distance D- 相对接近度C Relative
proximity C		 排序结果Sorting
result		 平均产量排序结果
Average yield
Sorting result
对照CK CK 2.618 1.350 0.340 15 15
甜菜碱
Betaine (T1)		 T1-1 3.111 0.882 0.221 17 17
T1-2 0.908 2.890 0.761 6 4
T1-3 1.079 2.705 0.715 7 7
脯氨酸
Proline (T2)		 T2-1 0.733 3.068 0.807 5 6
T2-2 0.642 3.168 0.832 4 5
T2-3 0.581 3.219 0.847 3 3
过氧化氢
Hydrogen
peroxide (T3)		 T3-1 3.659 0.377 0.093 18 18
T3-2 2.765 1.349 0.328 16 16
T3-3 2.340 1.624 0.410 13 13
苹果酸
Malic acid (T4)		 T4-1 2.236 1.741 0.438 12 12
T4-2 1.904 1.942 0.505 11 10
T4-3 2.505 1.493 0.373 14 14
氧化纳米锌
Nano zinc oxide (T5)		 T5-1 1.258 2.527 0.668 8 8
T5-2 0.335 3.496 0.913 2 2
T5-3 0.068 3.724 0.982 1 1
多胺
Polyamine (T6)		 T6-1 1.737 2.192 0.558 10 11
T6-2 1.445 2.365 0.621 9 9
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