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作物学报 ›› 2019, Vol. 45 ›› Issue (5): 728-739.doi: 10.3724/SP.J.1006.2019.83040

• 耕作栽培·生理生化 • 上一篇    下一篇

玉米-大豆带状种植中套作高光效玉米窄行穂位叶光合特性对弱光胁迫的响应

任永福,陈国鹏,蒲甜,陈诚,曾瑾汐,彭霄,马艳玮,杨文钰,王小春()   

  1. 四川农业大学农学院 / 农业部西南作物生理生态与耕作重点实验室 / 作物生理生态及栽培四川省重点实验室, 四川成都 611130
  • 收稿日期:2018-05-01 接受日期:2019-01-12 出版日期:2019-05-12 网络出版日期:2019-02-19
  • 通讯作者: 王小春
  • 基金资助:
    本研究由国家重点研发计划项目(2016YFD0300109);四川省育种攻关项目(2016NYZ0051-2);成都市农业技术成果应用示范项目(2015-NY01-00100-NC)

Responses of photosynthetic characteristics to low light stress in ear leaves of high photosynthetic efficiency maize at narrow row of maize-soybean strip intercropping system

Yong-Fu REN,Guo-Peng CHEN,Tian PU,Cheng CHEN,Jin-Xi ZENG,Xiao PENG,Yan-Wei MA,Wen-Yu YANG,Xiao-Chun WANG()   

  1. College of Agronomy, Sichuan Agricultural University / Key Laboratory of Crop Ecophysiology and Farming System in Southwest China, Ministry of Agriculture / Crop Ecophysiology and Cultivation Key Laboratory of Sichuan Province, Chengdu 611130, Sichuan, China
  • Received:2018-05-01 Accepted:2019-01-12 Published:2019-05-12 Published online:2019-02-19
  • Contact: Xiao-Chun WANG
  • Supported by:
    This study was supported by the National Key Research and Development Program of China(2016YFD0300109);the Breeding Research Project of Sichuan(2016NYZ0051-2);the Agricultural Technology Achievement Demonstration Project of Chengdu(2015-NY01-00100-NC)

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摘要：

玉米-大豆带状套作模式下, 宽窄行种植玉米, 窄行叶片存在着明显的光限制现象。采用低(A1: 众望玉18)、中(A2: 川单418)、高(A3: 荣玉1210)光效玉米品种, 玉米-大豆带状套作种植模式, 带宽固定为2 m, 每带种植2行玉米2行大豆, 设置大豆行距40 cm和玉米不同的窄行行距(B1: 20 cm; B2: 40 cm; B3: 单作)。探究套作高光效玉米窄行穂位叶在不同窄行光胁迫下的光合指标差异。结果表明, 与低光效和中光效品种玉米相比, 套作常规行距(40 cm)下, 高光效玉米品种(荣玉1210)窄行穗位叶光合速率、PEPCase活性分别显著高出46.9%、230.1%和11.8%、13.98%; 对弱光(10:00前、16:00后)的利用效率及叶绿体结构完整程度较高, 而叶绿素初始荧光(Fo)和最大荧光(Fm)较低; 随套作窄行行距减小, 3类品种玉米窄行穗位叶光合速率、光合作用关键酶活性均呈下降趋势, 叶绿素初始荧光(Fo)、最大荧光(Fm)及有效光化学量子产量(Fv'/Fm')均呈现不同程度的升高, 但均以套作高光效玉米变化幅度最小。套作高光效玉米在套作环境中窄行穗位叶的光合速率、光合作用关键酶活性、产量与单作差异均未达显著水平, 而低光效玉米和中光效玉米套作与单作相比, 光合速率、PEPCase活性显著降低28.9%、24.2%和7.4%、5.5%。因此, 不同玉米品种适应套作窄行光胁迫的能力差异显著, 套作高光效玉米(荣玉1210)在套作条件下仍具有相对理想的光合生理指标, 这为其适应套作光环境并获得高产提供了理论依据。

关键词: 玉米-大豆带状套作, 套作高光效, 穂位叶, 弱光胁迫, 光合特性

Abstract:

The photosynthesis of maize is inhibited in narrow row under maize-soybean relay strip intercropping system. In order to explore the internal mechanisms of photosynthesis characteristics responding to low light stress in ear leaf of intercropped maize cultivars with high photosynthetic efficiency, the low (A1: Zhongwangyu 18), middle (A2: Chuandan 418), and high (A3: Rongyu 1210) photosynthetic efficiency maize varieties were grown in maize-soybean strip intercropping system (two rows of maize intercropped with two rows of soybean) with different narrow spaces (B1: 20 cm; B2: 40 cm; B3: monoculture). The photosynthetic rate and PEPCase activity of “Rongyu 1210” were significantly higher than those of “Zhongwangyu 18” and “Chuandan 418”, respectively under the conventional row spacing (40 cm) and low light intensity conditions (before 10 a.m., after 4 p.m.). Additionally, “Rongyu 1210” had more complete chloroplast structure, which might result in lower initial fluorescence (Fo) and maximum fluorescence (Fm) of chlorophyll compared with “Zhongwangyu 18” and “Chuandan 418”. The Pn and activities of key enzymes decreased, but the initial fluorescence (Fo), maximum fluorescence (Fm) and effective photochemical quantum yield (Fv'/Fm') of chlorophyll increased with the decreasing narrow row spacing, Interestingly, the above indexes of “Rongyu 1210” were stable under the variable light environment. The photosynthetic rate, photosynthetic key enzyme activities, and yield of “Rongyu 1210” between monoculture and intercropping were not significantly different. However, the photosynthetic rate and PEPCase activity of “Zhongwangyu 18” and “Chuandan 418” in intercropping decreased by 28.9%, 24.2% and 7.4%, 5.5% compared with that in monoculture, respectively. In conclusion, the response of maize varieties to shade conditions in narrow row under intercropping system is different. Rongyu 1210, with a positive responding, that has better physiological indexes in photosynthesis than the other two maize varieties in the intercropping system. These results provide a theoretical explanation for excellent adaptability to the lower light environment and high yield in Rongyu 1210 under maize-soybean intercropping strip system.

Key words: maize-soybean strip intercropping, high photosynthetic efficiency, ear leaves, low light stress, photosynthetic characteristics

图1

田间种植及光分布示意图 MM代表单作。"

图2

不同处理对玉米窄行穂位叶光合作用参数的影响 MM代表单作。同一组数据中标以不同小写字母表示处理间在P < 0.05水平差异显著。"

图3

不同处理对玉米窄行穂位叶光合作用速率日变化的影响 MM代表单作。"

图4

不同处理对玉米窄行穂位叶光合作用关键酶的活性的影响 MM代表单作。同一组数据中标以不同小写字母表示处理间在P < 0.05水平差异显著。"

表1

不同处理对玉米窄行穂位叶叶绿素荧光参数的影响"

品种
Variety		 行距
Row
spacing		 Fo Fm Fv/Fm Fv′/Fm′
抽雄期
Tasseling stage		 灌浆期
Filling stage		 抽雄期
Tasseling stage		 灌浆期
Filling
stage		 抽雄期
Tasseling stage		 灌浆期
Filling stage		 抽雄期
Tasseling stage		 灌浆期
Filling stage
众望玉18 20 cm 3788.00 a 3438.00 b 19501.00 a 16201.00 a 0.76 c 0.77 c 0.13 a 0.14 b
Zhongwangyu 18 40 cm 2595.00 d 3370.00 c 13788.00 e 15819.00 b 0.78 ab 0.79 b 0.12 ab 0.13 bc
单作MM 2161.00 e 2600.00 e 12092.00 f 13566.00 d 0.79 a 0.81 a 0.10 c 0.12 c
川单418 20 cm 3527.00 b 3635.00 a 17138.00 b 15016.00 b 0.76 c 0.75 d 0.13 a 0.12 c
Chuandan 418 40 cm 3085.00 d 3326.00 c 16187.00 c 14527.00 c 0.77 b 0.76 cd 0.12 ab 0.12 c
单作MM 2345.00 e 2578.00 e 12284.00 f 12310.00 e 0.78 ab 0.79 b 0.12 ab 0.11 d
荣玉1210 20 cm 3481.00 b 2986.00 d 17400.00 b 14413.00 c 0.76 c 0.78 bc 0.11 bc 0.15 a
Rongyu 1210 40 cm 3272.00 c 2775.00 e 16816.00 c 13380.00 d 0.77 b 0.79 b 0.11 bc 0.14 b
单作MM 2964.00 d 1888.00 f 15585.00 d 10094.00 f 0.77 b 0.80 ab 0.10 c 0.12 c
众望玉18-avg. Zhongwangyu 18-avg. 2848.00 c 3136.00 a 15127.00 b 15195.33 a 0.78 a 0.79 a 0.12 a 0.13 a
川单418-avg. Chuandan 418-avg. 3052.33 b 3179.66 a 15203.00 b 13951.00 b 0.77 b 0.77 b 0.12 a 0.12 b
荣玉1210-avg. Rongyu 1210-avg. 3239.00 a 2549.66 b 16600.33 a 12629.00 b 0.77 b 0.79 a 0.11 b 0.13 a
20 cm-avg. 3598.66 a 3353.00 a 18013.00 a 15210.00 a 0.77 c 0.77 c 0.12 a 0.14 a
40 cm-avg. 3050.66 b 3157.00 b 15597.00 b 14575.33 b 0.78 b 0.78 b 0.12 a 0.13 b
MM-avg. 2490.00 c 2355.33 c 13320.33 c 11990.00 c 0.79 a 0.80 a 0.11 b 0.11 c
F值 F-value
品种 Varieties (V) 1.43* 2.02 0.99 2.36 5.21 4.14 2.23 4.02*
行距 Row spacing (RS) 4.07* 1.14 3.69 1.81 5.96* 0.88* 2.12 0.37*
品种×行距 V×RS 3.01* 1.50 2.61 1.65 4.70 2.18 2.17 1.83*

表2

不同处理对玉米窄行穂位叶叶绿体超微结构的影响"

品种
Variety		 行距
Row spacing		 叶绿体长度Chloroplast length (μm) 叶绿体宽度Chloroplast width (μm) 基粒厚度
Grains thickness (μm)		 淀粉颗粒数
Starch granule		 嗜锇颗粒数
Osmiophilic
particle
众望玉18 20 cm 7.12 bc 3.33 a 0.73 a 3.33 a 3.67 a
Zhongwangyu 18 40 cm 7.12 bc 3.10 b 0.72 ab 2.67 ab 3.33 a
单作MM 7.00 d 3.09 bc 0.69 b 2.00 b 1.33 bc
川单418 20 cm 7.13 b 3.26 ab 0.73 a 2.33 ab 2.33 b
Chuandan 418 40 cm 7.03 c 3.06 bc 0.68 bc 1.67 bc 1.33 bc
单作MM 6.98 e 3.06 bc 0.63 cd 1.33 bc 0.33 c
荣玉1210 20 cm 7.34 a 3.06 bc 0.65 c 1.00 c 1.00 cd
Rongyu 1210 40 cm 7.28 ab 2.99 d 0.61 d 0 d 0 d
单作MM 7.25 ab 2.98 d 0.61 d 0 d 0 d
F值 F-value
品种 Varieties (V) 55.28** 8.97* 16.44* 133.42** 16.66*
行距 Row pacing (RS) 11.48* 12.23* 7.18* 31.18** 8.79*
品种×行距 V×RS 0.07 0.11 0.95 0.59 9.09**

图5

透射电镜下的叶绿体超微结构 CHL: 叶绿体; GT: 基粒类囊体; CCHL: 紊乱的基粒类囊体; SG淀粉粒; OG: 嗜锇颗粒; GL: 基粒片层; SL: 基质片层。"

图6

不同处理对玉米产量的影响 MM代表单作。同一组数据中标以不同小写字母表示处理间在P < 0.05水平差异显著。"

表3

不同处理对玉米产量构成因素的影响"

品种
Variety		 行距
Row
spacing		 2015年 2016年
穗粒数
Grain number
per ear		 千粒重
1000-kernel
weight (g)		 穗数
Ear number
(×104 hm-2)		 穗粒数
Grain number
per ear		 千粒重
1000-kernel
weight (g)		 穗数
Ear number
(×104 hm-2)
众望玉18 20 cm 406.20 e 290.00 b 49764.30 b 429.12 d 324.04 a 50330.00 e
Zhongwangyu 18 40 cm 449.30 d 294.30 b 50357.10 b 420.80 d 316.20 b 54000.00 d
单作MM 507.20 b 289.10 b 57142.90 a 519.66 b 309.64 c 56000.00 c
川单418 20 cm 347.60 f 300.50 b 49521.40 b 459.47 c 263.06 e 54000.00 d
Chuandan 418 40 cm 468.70 c 292.60 b 57732.12 a 527.37 b 256.00 e 57330.00 b
单作MM 479.00 c 308.00 a 58992.90 a 588.85 a 255.46 e 59330.00 a
荣玉1210 20 cm 452.00 d 295.70 b 49764.30 b 394.54 e 294.60 d 56660.00 c
Rongyu 1210 40 cm 563.80 a 294.00 b 55512.25 ab 442.91 d 305.86 c 57660.00 b
单作MM 588.10 a 295.40 b 56664.30 a 472.68 c 310.20 c 57660.00 b
众望玉18-avg. Zhongwangyu 18-avg. 454.23 b 291.13 a 52421.43 b 456.52 b 316.62 a 53443.33 c
川单418-avg. Chuandan 418-avg. 431.76 b 300.36 a 55415.47 a 525.23 a 258.17 c 56886.66 b
荣玉1210-avg. Rongyu 1210-avg. 534.63 a 295.03 a 53980.28 a 436.71 c 303.55 b 57326.66 a
20 cm-avg. 401.93 c 295.40 a 49683.33 c 427.71 c 293.90 a 53663.33 c
40 cm-avg. 493.93 b 293.63 a 54533.82 b 463.69 b 292.68 a 56330.00 b
MM-avg. 524.76 a 297.50 a 57600.03 a 527.06 a 291.76 a 57663.33 a
F值 F-value
品种 Varieties (V) 25.78** 185.89** 1.32** 16.72** 66.00** 5.51**
行距 Row spacing (RS) 24.35** 3.49** 38.66* 27.18** 3.55 4.59*
品种×行距 V×RS 24.92** 76.45* 23.72** 1.63 2.00 0.72

表4

光合参数之间及其与产量的相关关系"

光合速率
Photosyn-
thetic rate		 气孔导度
Stomatal
conductance		 胞间CO2浓度
Intercellular CO2 concentration		 蒸腾速率
Transpiration rate		 穂粒数
Grain number per ear		 千粒重
1000-kernel weight		 穗数
Ear number
(×104 hm-2)		 产量
Yield
Photosynthetic rate 1.00 0.99** -0.80** 0.90** 0.54* 0.39 0.53* 0.62**
Stomatal conductance 1.00 -0.82** 0.92** 0.58* 0.37 0.54* 0.65**
Intercellular CO2 concentration 1.00 -0.97** -0.74** -0.24 -0.47 -0.73**
Transpiration rate 1.00 0.65** 0.27 0.51* 0.69**
Grain number per ear 1.00 -0.15 0.67** 0.95**
1000-kernel weight 1.00 0.18 0.05
Ear number (×104 hm-2) 1.00 0.85**
Yield 1.00
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