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作物学报

作物学报 ›› 2019, Vol. 45 ›› Issue (1): 118-130.doi: 10.3724/SP.J.1006.2019.84043

• 耕作栽培·生理生化 • 上一篇    下一篇

低温胁迫和赤霉素对花生种子萌发和幼苗生理响应的影响

常博文1,2(),钟鹏2,刘杰2,*(),唐中华3,高亚冰2,于洪久2,郭炜2   

  1. 1 黑龙江省农业科学院博士后科研工作站, 黑龙江哈尔滨 150001
    2 黑龙江省农业科学院农村能源研究所, 黑龙江哈尔滨 150001
    3 东北林业大学森林植物生态学教育部重点实验室, 黑龙江哈尔滨 150001
  • 收稿日期:2018-03-22 接受日期:2018-08-20 出版日期:2018-09-28 网络出版日期:2018-09-28
  • 通讯作者: 刘杰
  • 基金资助:
    本研究由黑龙江省农业科学院课题(2017BZ01);哈尔滨市科技创新人才研究专项和黑龙江省育繁推一体化项目资助(2017RAQYJ135)

Effect of low-temperature stress and gibberellin on seed germination and seedling physiological responses in peanut

Bo-Wen CHANG1,2(),Peng ZHONG2,Jie LIU2,*(),Zhong-Hua TANG3,Ya-Bing GAO2,Hong-Jiu YU2,Wei GUO2   

  1. 1 Postdoctoral Research Station, Heilongjiang Academy of Agricultural Sciences, Harbin 150001, Heilongjiang, China
    2 Rural Energy Institute, Heilongjiang Academy of Agricultural Science, Harbin 150001, Heilongjiang, China
    3 Key Laboratory of Forest Plant Ecology Ministry of Education, Northeast Forestry University, Harbin 150001, Heilongjiang, China
  • Received:2018-03-22 Accepted:2018-08-20 Published:2018-09-28 Published online:2018-09-28
  • Contact: Jie LIU
  • Supported by:
    This study was supported by the Program of Heilongjiang Academy of Agricultural Sciences(2017BZ01);Harbin Special Fund for Research of Innovative Talents of Science and Technology, and Heilongjiang Integration Program of “Breeding, Reproduction and Pushing”.(2017RAQYJ135)

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摘要：

低温是影响我国广大花生产区春花生发芽的主要因素之一。本文以不同生态区的30个花生品种为实验材料, 研究了倒春寒天气诱导的低温胁迫对花生出苗的影响, 以出苗率为标准筛选出4个耐低温的花生品种(阜花17、阜花12、冀花16、冀花18)和4个不耐低温的品种(鲁花11、白沙1016、正农黑花生1号、白玉)于温室测定了4℃低温和赤霉素(GA3)处理后种子发芽相关指标和幼苗生理指标。结果表明, 4℃对耐低温花生品种发芽率、发芽指数影响不显著, 但种子活力指数和芽长呈现下降趋势; 4℃处理后, 不耐低温品种幼苗相对膜透性和MDA含量上升幅度更高, 耐低温品种幼苗的可溶性糖和游离脯氨酸含量上升幅度更大。GA3显著促进4℃低温处理后花生种子萌发和种子活力, 抑制了花生幼苗在低温处理后相对膜透性和丙二醛的上升, 提高了可溶性糖、可溶性蛋白、游离脯氨酸含量。研究表明, 赤霉素促进低温胁迫下种子萌发和幼苗生长的最佳浓度是300 μmol L -1。发芽率与相对膜透性和丙二醛含量显著负相关, 与可溶性糖、脯氨酸含量显著正相关。温度影响发芽率的品种间差异较大, 常温下赤霉素对不同耐低温品种的发芽率影响较小。本研究为耐低温花生种质资源创新和新品种培育提供了理论依据, 为研究赤霉素对不同花生品种耐低温性影响的生理机制提供了基础。

关键词: 花生, 倒春寒, 低温胁迫, 赤霉素, 种子萌发, 幼苗生理响应

Abstract:

From 30 peanut cultivars, we screened four cultivars with strong low-temperature-resistance (Fuhua 17, Fuhua 12, Jihua 16, Jihua 18) and four cultivars with weak low-temperature-resistance (Luhua 11, Baisha 1016, Zhengnongheihuasheng 1, Baiyu), and measured related indicators of seed germination and physiological indexes of seedlings under low temperature (4oC) and GA3 treatments. There was no significant difference in rate of emergence and germination index of strong low-temperature- resistant cultivars with or without 4oC treatments, but decrease in hypocotyl length and seed vigor under low temperature. The relative membrane permeability and MDA content of four cultivars with weak low-temperature-resistance had higher ascensional range. The contents of soluble sugar, soluble protein and free proline of cultivars with strong low-temperature resistance had smaller reduction. GA3 facilitated the rate of emergence and seed vigor of all peanut seeds, as well as promoted the contents of soluble sugar, soluble protein and free proline of seedlings, but suppressed the uptrend of relative membrane permeability and MDA content of peanut seedlings under low temperature. The best concentration of GA3 promoting seed germination and seedling growth of peanut with low temperature treatment is 300 μmol L -1. The rate of emergence had significantly negative correlation with relative membrane permeability and MDA, and obviously positive correlation with the contents of soluble sugar or free proline. The temperature had greater influence on germination rate, but the effect of gibberellin on difference of germination rate between different varieties was smaller. This study provided a theoretical basis for germplasm resources innovation and breeding new peanut cultivars with strong low-temperature resistance, as well as for studying the physiological mechanism of gibberellin on chilling tolerance of different peanut varieties.

Key words: peanut, late spring chilling, low-temperature stress, gibberellin, seed germination, seedling physiological responses

表1

黑龙江省倒春寒等级指标"

倒春寒等级
Late spring chilling grades		 最大降温幅度ΔT
Maximum cooling rate (oC)		 气温距平(δT)
Temperature departure (oC)		 持续时间
Consecutive time (d)
轻微 Light 3.0 < ΔT24 ≤ 5.0 or 5.0 < ΔT48 ≤ 8.0 -2.0 ≤ δT < -1.0 ≥ 3
中度偏轻 Moderate partial light 5.0 < Δ.0erate or 8.0 < Δ.0erate -3.0 ≤ δT < -2.0 ≥ 3
中度 Moderate 8.0 < ΔT24 ≤ 10.0 or 10.0 < ΔT48 ≤ 12.0 -5.0 ≤ δT < -3.0 ≥ 5
严重 Severe 10.0 < ΔT24 or 12.0 < ΔT48 δT < -5.0 ≥ 5

图1

2017年5月至6月气温变化 MAX T: 日最高气温; MIN T: 日最低气温; Average: 日平均气温。"

表2

播种1个月后各花生品种出苗率"

品种
Cultivar		 出苗率
Seedling emergence rate (%)		 标准误差
Standard error		 差异显著性
Sig. 5%
白玉 Baiyu 45.51 8.43 a
正农黑花生1号 Zhengnongheihuasheng 1 49.79 10.08 ab
白沙1016 Baisha 1016 50.28 8.52 ab
七彩花生 Qicaihuasheng 58.63 3.89 bc
鲁花11 Luhua 11 59.63 9.76 bcd
花育22 Huayu 22 67.33 10.25 cde
荣丰8号 Rongfeng 8 68.42 9.58 cde
丰花5号 Fenghua 5 69.29 5.32 de
丰花3号 Fenghua 3 72.75 5.03 ef
冀花11 Jihua 11 72.99 6.34 ef
丰花1号 Fenghua 1 73.30 4.82 efg
海花1号 Haihua 1 73.54 3.22 efg
中农108 Zhongnong 108 74.00 8.92 efg
山花10号 Shanhua 10 74.33 3.24 efg
中花800 Zhonghua 800 75.00 4.77 efg
拔二罐 Baerguan 76.38 6.64 efgh
花育23 Huayu 23 81.73 4.38 fghi
品种
Cultivar		 出苗率
Seedling emergence rate (%)		 标准误差
Standard error		 差异显著性
Sig. 5%
冀花4号 Jihua 4 82.06 4.22 fghi
冀花9号 Jihua 9 82.63 1.04 fghij
冀花8号 Jihua 8 84.22 4.13 ghijk
寿花820 Shouhua 820 86.38 5.66 hijkl
花育36 Huayu 36 89.71 4.09 ijkl
四粒红 Silihong 90.19 5.24 ijkl
冀花10号 Jihua 10 90.95 2.74 ijkl
花育20 Huayu 20 91.43 2.74 ijkl
冀花324 Jihua 324 93.40 1.89 jkl
冀花18 Jihua 18 94.56 4.16 kl
冀花16 Jihua 16 95.27 1.49 l
阜花12 Fuhua 12 95.36 3.27 l
阜花17 Fuhua 17 97.22 1.26 l

图2

不同温度对花生种子发芽率的影响 FH17: 阜花17; FH12: 阜花12; JH16: 冀花16; JH18: 冀花18; BS1016: 白沙1016; LH11: 鲁花11; HHS: 正农黑花1号; BHS: 白玉花生。图柱上不同字母表示处理之间的0.05水平差异显著。"

表3

低温胁迫和GA3对花生种子发芽率、发芽指数、芽长和种子活力的影响"

花生品种
Peanut cultivar		 对照组
Control group		 处理组 Treatment group
4℃ 4℃+100
μmol L-1 GA3		 4℃+200
μmol L-1 GA3		 4℃+300
μmolL-1 GA3
发芽率Seed germination rate (%)
阜花17 Fuhua 17 98.00±1.27 ab 87.33±1.73 b 89.33±1.04 b 91.67±0.99 b 95.33±1.46 b
阜花12 Fuhua 12 97.33±1.44 ab 85.33±1.02 b 89.33±2.13 b 92.33±0.57 b 94.00±0.98 b
冀花16 Jihua 16 98.67±0.67 ab 85.00±1.67 b 90.33±2.06 b 92.33±1.33 b 94.00±1.04 b
冀花18 Jihua 18 99.00±1.09 b 84.67±2.56 b 90.00±2.01 b 93.67±1.77 b 95.00±2.76 b
白沙1016 Baisha 1016 99.00±0.78 b 48.33±4.37 a 51.67±3.22 a 57.00±4.06 a 60.67±1.34 a
鲁花11 Luhua 11 96.33±1.78 a 50.67±5.34 a 54.00±2.35 a 60.67±3.33 a 65.33±3.32 a
正农黑花生1号 Zhengnongheihuasheng 1 99.00±0.33 b 48.67±4.43 a 54.00±4.78 a 56.00±5.49 a 60.67±3.97 a
白玉Baiyu 98.67±1.08 ab 45.67±4.56 a 50.00±1.98 a 54.33±3.46 a 61.67±3.22 a
芽长Hypocotyl length (cm)
阜花17 Fuhua 17 5.36±0.46 a 4.08±0.55 d 4.23±0.33 c 4.48±0.18 d 4.94±0.25 c
阜花12 Fuhua 12 4.98±0.45 a 3.55±0.34 cd 3.76±0.26 c 4.01±0.18 c 4.66±0.21 c
冀花16 Jihua 16 5.23±0.22 a 3.46±0.11 cd 3.99±0.32 c 4.31±0.15 cd 4.79±0.54 c
冀花18 Jihua 18 5.06±0.33 a 3.33±0.21 c 3.79±0.14 c 4.29±0.21 cd 4.48±0.28 c
白沙1016 Baisha 1016 5.16±0.42 a 1.88±0.09 a 2.06±0.10 a 2.14±0.13 a 2.36±0.18 a
鲁花11 Luhua 11 4.92±0.46 a 2.19±0.08 b 2.36±0.09 b 2.65±0.12 b 2.95±0.15 b
正农黑花生1号 Zhengnongheihuasheng 1 4.86±0.50 a 2.21±0.09 b 2.37±0.21 b 2.56±0.13 b 2.83±0.14 b
白玉 Baiyu 5.26±0.45 a 2.06±0.12 ab 2.24±0.21 ab 2.41±0.19 ab 2.61±0.25 ab
发芽指数 Germination index
阜花17 Fuhua 17 21.34±2.14 a 16.24±1.24 b 17.89±2.45 b 18.56±1.11 b 20.55±2.01 b
阜花12 Fuhua 12 20.16±1.99 a 15.06±1.45 b 16.63±1.84 b 17.94±1.86 b 19.34±1.08 b
冀花16 Jihua 16 21.33±2.01 a 16.12±1.33 b 17.64±1.04 b 18.48±1.75 b 20.26±1.88 b
冀花18 Jihua 18 22.21±1.99 a 14.90±1.23 b 15.78±1.64 b 17.30±1.13 b 19.10±1.86 b
白沙1016 Baisha 1016 20.55±2.01 a 9.00±1.08 a 10.35±1.44 a 11.56±1.07 a 12.62±2.33 a
鲁花11 Luhua 11 19.17±2.88 a 10.72±1.64 a 11.69±1.37 a 13.16±1.05 a 14.97±1.75 a
正农黑花生1号 Zhengnongheihuasheng 1 20.22±2.54 a 9.90±1.85 a 11.00±1.32 a 12.02±1.56 a 13.34±1.33 a
白玉 Baiyu 19.31±2.04 a 9.38±1.15 a 10.94±1.46 a 12.26±1.35 a 13.15±1.78 a
种子活力指数 Seed vigor
阜花17 Fuhua 17 114.38±10.11 b 66.28±6.91 c 75.67±5.90 d 83.17±7.45 c 101.52±8.99 c
阜花12 Fuhua 12 100.38±8.45 ab 53.46±6.23 bc 62.53±5.34 c 71.94±7.89 c 90.12±10.11 c
冀花16 Jihua 16 111.58±10.58 ab 55.78±6.03 bc 70.38±5.84 cd 79.65±7.77 c 97.05±8.09 c
冀花18 Jihua 18 112.39±8.23 b 49.61±5.89 b 59.81±7.82 c 74.21±8.19 c 85.56±8.74 c
白沙1016 Baisha 1016 106.04±11.56 ab 16.92±3.65 a 21.33±2.98 a 24.74±3.02 a 29.78±2.00 a
鲁花11 Luhua11 93.15±10.67 a 23.68±3.98 a 27.70±2.76 b 33.69±3.14 b 42.35±4.23 b
正农黑花生1号 Zhengnongheihuasheng 1 99.46±11.49 ab 21.67±2.89 a 25.97±3.01 ab 31.85±2.76 b 39.35±3.45 b
白玉 Baiyu 101.59±10.97 ab 19.32±2.05 a 24.50±2.86 ab 29.55±3.56 ab 34.33±3.89 ab

表4

低温胁迫和GA3对花生幼苗相对膜透性、丙二醛、可溶性糖、可溶性蛋白和游离脯氨酸含量的影响"

生理指标
Physical
indexes		 花生品种
Peanut cultivar		 对照组
Control group		 处理组Treatment group
4℃ 4℃+100
μmol L-1 GA3		 4℃+200
μmol L-1 GA3		 4℃+300
μmol L-1 GA3
相对膜透性
Relative membrane permeability (%)		 阜花17 Fuhua 17 17.10±1.89 ab 24.63±2.08 a 21.63±1.65 a 20.33±1.32 a 18.71±1.16 a
阜花12 Fuhua 12 18.50±1.92 b 26.98±1.36 a 24.56±1.01 b 22.68±1.06 ab 20.22±1.45 ab
冀花16 Jihua16 15.92±1.65 a 26.88±2.02 a 25.69±1.54 b 23.79±0.98 b 21.25±0.99 b
冀花18 Jihua 18 15.32±1.54 a 25.54±1.85 a 24.46±1.99 ab 23.03±1.33 b 21.46±1.25 b
白沙1016 Baisha 1016 16.43±1.98 ab 31.23±1.89 b 30.36±1.45 c 28.46±1.85 c 26.18±1.28 c
鲁花11 Luhua 11 17.45±2.01 ab 31.23±1.97 b 30.56±1.67 c 28.12±1.11 c 26.54±1.98 c
正农黑花生1号
Zhengnongheihuasheng 1		 17.95±1.85 ab
 30.91±1.16 b
 28.63±1.22 c
 27.56±1.00 c
 25.87±1.18 c
白玉Baiyu 15.99±1.89 a 30.88±1.96 b 29.36±1.02 c 27.98±1.45 c 26.87±1.08 c
丙二醛含量
MDA content
(μmol g-1 FW-1)		 阜花17 Fuhua 17 7.13±0.95 a 9.56±1.04 a 8.66±0.83 a 7.84±0.66 a 7.16±0.58 a
阜花12 Fuhua 12 7.56±0.96 a 9.84±0.68 a 8.92±0.51 a 8.06±0.53 a 7.98±0.73 a
冀花16 Jihua 16 6.35±0.83 a 10.19±1.01 a 9.13±0.77 a 8.46±0.49 a 7.39±0.50 a
冀花18 Jihua 18 6.84±0.77 a 10.23±0.93 a 9.04±1.00 a 8.16±0.67 a 7.28±0.63 a
白沙1016 Baisha 1016 6.45±0.99 a 19.03±0.95 b 17.85±0.73 b 16.32±0.93 c 14.88±1.06 c
鲁花11 Luhua 11 7.26±1.01 a 18.32±0.99 b 16.98±0.84 b 15.36±0.56 bc 13.86±0.99 bc
正农黑花生1号
Zhengnongheihuasheng 1		 7.06±0.93 a
 19.26±0.58 b
 16.54±1.05 b
 14.46±0.50 b
 11.65±1.46 b
白玉 Baiyu 6.98±0.95 a 19.69±0.98 b 17.05±1.51 b 15.24±0.73 bc 12.74±1.22 bc
可溶性糖
Soluble sugar content (mg g-1 FW-1)		 阜花17 Fuhua 17 7.46±1.95 b 18.75±1.49 b 21.36±0.83 b 24.68±1.66 bc 29.00±1.28 b
阜花12 Fuhua 12 7.96±1.36 b 18.32±1.38 b 20.69±0.51 b 24.12±1.53 bc 28.56±1.73 b
冀花16 Jihua 16 8.63±1.26 b 17.11±1.51 b 21.63±0.77 b 25.55±1.49 c 27.36±1.00 b
冀花18 Jihua 18 7.78±1.53 b 18.56±1.43 b 20.45±1.00 b 22.69±1.67 b 28.33±1.63 b
白沙1016 Baisha 1016 5.55±0.99 ab 8.78±1.95 a 10.96±0.73 a 12.15±1.90 a 14.15±1.56 a
鲁花11 Luhua 11 4.99±1.01 a 8.82±1.49 a 10.65±0.84 a 12.21±1.56 a 14.54±0.99 a
正农黑花生1号
Zhengnongheihuasheng 1		 5.87±0.93 ab
 9.74±1.28 a
 11.96±1.05 a
 13.33±1.51 a
 15.78±1.46 a
白玉Baiyu 6.45±1.34 ab 8.64±1.48 a 11.62±1.51 a 13.05±1.73 a 14.51±1.22 a
生理指标
Physical
indexes		 花生品种
Peanut cultivar		 对照组
Control group		 处理组Treatment group
4℃ 4℃+100
μmol L-1 GA3		 4℃+200
μmol L-1 GA3		 4℃+300
μmol L-1 GA3
可溶性蛋白
Soluble protein content (μmol g-1 FW-1)		 阜花17 Fuhua 17 16.49±2.09 a 25.45±0.98 b 26.85±1.65 b 27.96±1.66 b 29.64±1.28 b
阜花12 Fuhua 12 17.68±1.22 a 23.78±0.76 ab 26.45±1.01 c 28.65±1.53 b 30.77±1.73 b
冀花16 Jihua 16 17.45±2.52 a 24.36±1.22 ab 25.85±1.54 bc 26.56±1.49 ab 29.86±1.00 b
冀花18 Jihua 18 16.12±2.06 a 25.88±1.85b 26.74±1.99 b 27.65±1.67 b 30.55±1.03 b
白沙1016 Baisha 1016 18.02±1.98 a 23.71±1.89 ab 24.56±1.45 ab 25.74±1.23 ab 26.16±1.56 a
鲁花11 Luhua 11 17.68±2.01 a 23.52±1.67 ab 23.65±1.67 ab 24.31±1.56 a 26.99±0.99 a
正农黑花生1号
Zhengnongheihuasheng 1		 16.59±1.85 a
 22.34±1.56 a
 23.99±1.35 ab
 24.15±1.50 a
 26.59±1.46 a
白玉Baiyu 17.66±2.68 a 21.90±1.96 a 23.06±1.02 a 23.94±1.73 a 25.35±1.22 a
游离脯氨酸
Free proline
(μmol g-1 FW-1)		 阜花17 Fuhua 17 60.01±7.23 a 83.95±7.24 a 100.65±6.23 ab 112.14±8.66 ab 135.63±7.85 bc
阜花12 Fuhua 12 65.99±7.01 a 85.45±8.36 a 99.46±7.12 ab 116.32±8.12 b 140.63±7.78 c
冀花16 Jihua 16 61.23±8.96 a 86.56±8.11 a 101.25±7.52 ab 110.36±7.78 ab 136.44±8.33 bc
冀花18 Jihua 18 59.46±8.25 a 80.11±8.46 a 103.66±6.66 b 120.56±8.36 b 133.78±7.55 b
白沙1016 Baisha 1016 63.19±4.48 a 78.25±7.12 a 90.26±7.16 a 108.36±7.45 ab 122.32±6.06 ab
鲁花11 Luhua 11 61.85±6.85 a 79.33±7.56 a 90.29±8.56 a 104.85±6.21 a 118.62±7.60 a
正农黑花生1号
Zhengnongheihuasheng 1		 58.36±6.99 a
 74.63±7.98 a
 91.29±7.31 a
 102.24±6.65 a
 121.55±8.32 ab
白玉 Baiyu 62.45±7.11 a 79.35±8.24 a 95.11±8.56 ab 104.15±7.12 a 119.62±8.41 a

表5

种子萌发相关指标与幼苗生理生化指标的相关系数"

GR GI SV HL RMP MDA SS SP FP
发芽率GR 1
发芽指数GI 0.935** 1
种子活力指数SV 0.923** 0.982** 1
芽长HL 0.961** 0.936** 0.964** 1
相对膜透性RMP -0.923** -0.888** -0.938** -0.981** 1
MDA含量 MDA -0.997** -0.934** -0.934** -0.972** 0.943** 1
可溶性糖含量SS 0.973** 0.872** 0.897** 0.970** -0.960** -0.983** 1
可溶性蛋白含量SP 0.742* 0.556 0.583 0.737* -0.718* -0.742* 0.829* 1
游离脯氨酸含量FP 0.864** 0.882** 0.810* 0.820* -0.745* -0.834* 0.779* 0.64 1

表6

效应因子正态分布检验"

A B C Shapiro-Wilk检验 Shapiro-Wilk test
统计量Statistic 自由度df 显著性Sig.
耐低温品种
Low-temperature tolerant
varieties (A1)		 对照Control 0 0.966 4 0.814
300 μmol L-1 0.873 4 0.310
4℃ 0 0.876 4 0.322
300 μmol L-1 0.729 4 0.024
不耐低温品种
Low-temperature susceptible varieties (A2)		 对照Control 0 0.827 4 0.161
300 μmol L-1 0.917 4 0.519
4℃ 0 0.965 4 0.811
300 μmol L-1 0.791 4 0.086

表7

不同效应因子交互效应方差分析结果"

效应因子
Source		 显著性
Sig.
A 0.000
B 0.000
C 0.000
A×B 0.000
A×C 0.011
B×C 0.000
A×B×C 0.011

表8

效应因子交互作用简单效应检验"

效应因子
Source		 F值
F-value		 显著性
Sig.
B WITHIN A1 3.23 0.084
B WITHIN A2 77.70 0.000
C WITHIN A1 1.19 0.286
C WITHIN A2 2.52 0.124
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