施氮肥缓解臭氧对小麦光合作用和产量的影响

doi:10.3724/SP.J.1258.2011.00523

摘要/Abstract

摘要：

以小麦(Triticum aestivum)品种‘扬麦16’为试材, 利用开放式空气臭氧(O₃)浓度升高平台, 研究了增施氮(N)肥对O₃对小麦光合作用和产量影响的缓解作用。结果表明, O₃胁迫下灌浆期小麦的净光合速率(P_n)、气孔导度(G_s)、蒸腾速率(T_r)、叶绿素a (Chl a)、叶绿素b (Chl b)、类胡萝卜素(Car)、总叶绿素含量(Chl t)和可溶性蛋白的含量显著降低, 降幅分别为28.95%、31.79%、23.17%、58.89%、68.64%、22.89%、60.31%和32.00%; 胞间CO₂浓度(C_i)变化很小; 成熟期生物量和收获时产量也明显下降, 降幅分别为12.23%和12.63%; 而增施N肥可以增加小麦灌浆期的P_n、Chl a、Chl b、可溶性蛋白的含量, 进而增加小麦生物量和产量, 增幅分别为25.66%、83.05%、121.57%、30.33%、14.94%和10.67%, 而对C_i、G_s、T_r、Car含量无明显影响。O₃和N肥对小麦叶片的P_n、Chl t及可溶性蛋白含量有明显的交互作用。因此, 在大气O₃浓度升高条件下增施N肥对小麦O₃损伤有一定的缓解作用。

关键词: 生物量, 缓解, 氮肥, 臭氧, 光合作用, 产量

Abstract:

Aims Our objective was to study the interactive influences of ozone (O₃) and nitrogen (N) on photosynthesis and yield in winter wheat (Triticum aestivum).

Methods The winter wheat was exposed to two levels of O₃ (ambient and 1.5 ambient) and two levels of N supply (210 and 250 kg·hm^-2) under field conditions.

Important findings O₃ exposure significantly reduced the net photosynthetic rate (P_n), stomatal conductance (G_s) and transpiration rate (T_r) of wheat leaves in the filling stage by 28.95%, 31.79 % and 23.17%, respectively. O₃ exposure also significantly reduced the content of chlorophyll a (Chl a), chlorophyll b (Chl b), carotene (Car), total chlorophyll (Chl t) and soluble protein in the filling stage by 58.89%, 68.64%, 22.89%, 60.31% and 32.00%, respectively, while intercellular CO₂ concentration (C_i) changed slightly. Biomass in the maturing stage and yield of the wheat were also reduced by elevated O₃ by 12.23% and 12.63%, respectively. High N availability significantly increased P_n, Chl a, Chl b, soluble protein, biomass and yield of the wheat leaves by 25.66%, 83.05%, 121.57%, 30.33%, 14.94% and 10.67%, respectively, while G_s, C_i, T_r and Car were influenced slightly by high N, which indicated that the increase of P_n was mainly caused by the increment of Chl a, Chl b and soluble protein. The interactive effects of O₃ and N on the concentrations of P_n, Chl a, Chl b and soluble protein were significant. These results suggest that sufficient N supply can modify the effects of elevated O₃ on photosynthesis and yield in wheat.

Key words: biomass, mediation, nitrogen, ozone, photosynthesis, yield

陈娟, 曾青, 朱建国, 刘钢, 曹际玲, 谢祖彬, 唐昊冶, 小林和彦. 施氮肥缓解臭氧对小麦光合作用和产量的影响. 植物生态学报, 2011, 35(5): 523-530. DOI: 10.3724/SP.J.1258.2011.00523

CHEN Juan, ZENG Qing, ZHU Jian-Guo, LIU Gang, CAO Ji-Ling, XIE Zu-Bin, TANG Hao-Ye, KAZUHIKO Kobayashi. Nitrogen supply mitigates the effects of elevated [O₃] on photosynthesis and yield in wheat. Chinese Journal of Plant Ecology, 2011, 35(5): 523-530. DOI: 10.3724/SP.J.1258.2011.00523

导出引用管理器 EndNote|Ris|BibTeX

链接本文: https://www.plant-ecology.com/CN/10.3724/SP.J.1258.2011.00523

https://www.plant-ecology.com/CN/Y2011/V35/I5/523

图/表 4

图1 O3和N交互作用对小麦叶片净光合速率、气孔导度、胞间CO2浓度和蒸腾速率的影响(平均值±标准偏差)。AHN, 对照处理高N水平; ANN, 对照处理常规N水平; FNN, O3处理常规N水平。FHN, O3处理高N水平。不同字母表示差异显著(p < 0.05)。

Fig. 1 Interactive effects of elevated O3 and nitrogen on net photosynthesis rate (Pn), stomatic conductance (Gs), intercellular CO2 concentration (Ci) and transpiration rate (Tr) of the wheat leaves (mean ± SD). ANN, ambient ozone and normal nitrogen. AHN, ambient ozone and high nitrogen; FNN, elevated ozone and normal nitrogen; FHN, elevated ozone and high nitrogen. Different letters mean significance at p < 0.05 level.

图2 O3和N交互作用对小麦叶片光合色素含量的影响(平均值±标准偏差)。AHN, 对照处理高N水平; ANN, 对照处理常规N水平; FHN, O3处理高N水平; FNN, O3处理常规N水平。不同字母表示差异显著(p < 0.05)。

Fig. 2 Interactive effects of elevated O3 and nitrogen on the photosynthetic pigment content of wheat leaves (mean ± SD). ANN, ambient ozone and normal nitrogen; AHN, ambient ozone and high nitrogen; FNN, elevated ozone and normal nitrogen; FHN, elevated ozone and high nitrogen. Different letters mean significance at p < 0.05 level.

图3 O3和N交互对小麦叶片可溶性蛋白含量的影响(平均值±标准偏差)。ANN, 对照处理常规N水平; AHN, 对照处理高N水平; FNN, O3处理常规N水平; FHN, O3处理高N水平。不同字母表示差异显著(p < 0.05)。

Fig. 3 Interactive effects of elevated O3 and nitrogen on soluble protein content in wheat leaves (mean ± SD). ANN, ambient ozone and normal nitrogen; AHN, ambient ozone and high nitrogen; FNN, elevated ozone and normal nitrogen; FHN, elevated ozone and high nitrogen. Different letters mean significance at p < 0.05 level.

表1 O3和N肥交互作用对小麦产量和不同生育期生物量(kg·hm-2)的影响(平均值±标准偏差)

Table 1 Interactive effects of elevated O3 and nitrogen on the yield and biomass of different growth period (kg·hm-2) of wheat (mean ± SD)

处理 Treatment		总生物量 Total biomass			产量 Yield
处理 Treatment		抽穗期 Heading	开花期 Blooming	成熟期 Maturing	收获期 Harvesting
NN	A	9 003 ± 897	12 195 ± 1235	14 657 ± 140	8 029 ± 396
	F	10 565 ± 1 023	11 700 ± 676	12 865 ± 428	7 014 ± 259
HN	A	9 809 ± 690	12 252 ± 1 310	14 491 ± 1 031	8 224 ± 548
	F	11 324 ± 814	11 831 ± 322	14 787 ± 447	7 762 ± 265
N		ns	ns	ns	ns
O₃		ns	ns	ns	*
N × O₃		ns	ns	ns	ns

参考文献 27

[1]	Andrews JT, Lorimer GH (1987). Rubisco: structure, mechanisms, and prospects for improvement. In: Stumpf PK, Conn EF eds. The Biochemistry of Plants: A Comprehensive Treatise, Photosynthesis. Academic Press, San Diego. 10, 131-218.
[2]	Ashmore MR (2005). Assessing the future global impacts of ozone on vegetation. Plant, Cell & Environment, 28, 949-964. DOI URL
[3]	Cao JL (曹际玲), Wang L (王亮), Zeng Q (曾青), Liang J (梁晶), Tang HY (唐昊冶), Xie ZB (谢祖彬), Liu G (刘钢), Zhu JG (朱建国), Kobayashi K (2009). Characteristics of photosynthesis in wheat cultivars with different sensitivity to ozone under O₃-free air control enrichment conditions. Acta Agronomica Sinica (作物学报), 35, 1500-1507. (in Chinese with English abstract)
[4]	Chen Z, Wang XK, Feng ZZ, Zheng FX, Duan XN, Yang WR (2008). Effects of elevated ozone on growth and yield of field-grown rice in Yangtze River Delta. Journal of Environmental Sciences, 20, 320-325. DOI URL
[5]	Feng ZZ, Pang J, Kobayashi K, Zhu JG (2011). Differential responses in two varieties of winter wheat to elevated ozone concentration under fully open-air field conditions. Global Change Biology, 17, 580-591.
[6]	Häikiö E, Freiwald V, Silfver T, Beuker E, Holopainen T, Oksanen E (2007). Impacts of elevated ozone and nitrogen on growth and photosynthesis of European aspen (Populus tremula) and hybrid aspen (P. tremula × Populus tremuloides) clones. Canadian Journal of Forest Research, 37, 2326-2336.
[7]	Handley T, Grulke NE (2008). Interactive effects of O₃ exposure on California black oak (Quercus kelloggii Newb.) seedlings with and without N amendment. Environmental Pollution, 156, 53-60. DOI URL PMID
[8]	Kobayashi K, Okada M (1995). Effects of ozone on the light use of rice (Oryza sativa L.). Plants Agriculture, Ecosystems and Environment, 53, 1-12.
[9]	Long SP, Naidu SL (2002). Effects of oxidants at the biochemical, cell and physiological levels, with particular reference to ozone. In: Bell JNB, Treshow M eds. Air Pollution and Plants 2nd edn. John Wiley and Sons Ltd., West Sussex. 69-88.
[10]	Makino A, Mae T, Ohira K (1983). Photosynthesis and ribulose-1, 5-bisphosphate carboxylase in rice leaves. Plant Physiology, 73, 1002-1007. URL PMID
[11]	Makino A, Mae T, Ohira K (1986). Colorimetric measurement of protein stained with Coomassie Brilliant Blue R on sodium dodecyl sulfate-polyacrylamide gel electrophoresis by eluting with formamide. Agricultural and Biology Chemistry, 50, 1911-1912.
[12]	Maurer S, Matyssek R, Günthardt-Goerg MS, Landolt W, Einig W (1997). Nutrition and the ozone sensitivity of birch (Betula pendula). I. Responses at the leaf level. Trees, 12, 1-10.
[13]	Morgan PB, Mies TA, Bollero GA, Nelson RL, Long SP (2006). Season-long elevation of ozone concentration to projected 2050 levels under fully open-air conditions substantially decreases the growth and production of soybean. New Phytologist, 170, 333-343. URL PMID
[14]	Nakaji T, Izuta T (2001). Effects of ozone and/or excess soil nitrogen on growth, needle gas exchange rates and rubisco contents of Pinus densiflora seedlings. Water, Air, and Soil Pollution, 130, 971-976.
[15]	Nakaji T, Kobayashi T, Kuroha M, Omori K, Matsumoto Y, Yonekura T, Watanabe K, Utriainen J, Izuta T (2004). Growth and nitrogen availability of red pine seedlings under high nitrogen load and elevated ozone. Water, Air, and Soil Pollution, 4, 277-287.
[16]	Pang J, Kobayashi K, Zhu JG (2009). Yield and photosynthetic characteristics of flag leaves in Chinese rice (Oryza sativa L.) varieties subjected to free-air release of ozone. Agriculture, Ecosystems and Environment, 132, 203-211.
[17]	Singh P, Agrawal M, Agrawal SB (2009). Evaluation of physiological, growth and yield responses of a tropical oil crop (Brassica campestris L. var. kranti) under ambient ozone pollution at varying NPK levels. Environmental Pollution, 157, 871-880. DOI URL PMID
[18]	Sitch S, Cox PM, Collins WJ, Huntingford C (2007). Indirect radiative forcing of climate change through ozone effects on the land-carbon sink. Nature, 448, 791-795. DOI URL PMID
[19]	Utriainen J, Holopainen T (2001a). Influence of nitrogen and phosphorus availability and ozone stress on Norway spruce seedlings. Tree Physiology, 21, 447-456. DOI URL PMID
[20]	Utriainen J, Holopainen T (2001b). Nitrogen availability modifies the ozone responses of Scots pine seedlings exposed in an open-field system. Tree Physiology, 21, 1205-1213. URL PMID
[21]	Vilhena JC, Barnes J (2001). Does nitrogen supply affect the response of wheat (Triticum aestivum cv. Hanno) to the combination of elevated CO₂ and O₃? Journal of Experimental Botany, 52, 1901-1911. URL PMID
[22]	Wang L (王亮), Zeng Q (曾青), Feng ZZ (冯兆忠), Zhu JG (朱建国), Tang HY (唐昊冶), Chen X (陈曦), Xie ZB (谢祖彬), Liu G (刘钢), Kobayashi K (2009). Photosynthetic damage induced by elevated O₃ in two varieties of winter wheat with free air controlled enrichment approach. Environmental Science (环境科学), 30, 527-534. (in Chinese with English abstract)
[23]	Watanabe M, Yamaguchi M, Matsumura H, Kohno Y, Izuta T (2008). Effects of ozone on the growth and photosynthesis of Castanopsis sieboldii seedlings growth under different nitrogen loads. Journal of Agricultural Meteorology, 64, 143-155.
[24]	Yamaguchi M, Watanabe M, Matsuo N, Naba J, Funada R, Fukami M, Matsumura H, Kohno Y, Izuta T (2007). Effects of nitrogen supply on the sensitivity to O₃ of growth and photosynthesis of Japanese beech (Fagus crenata) seedlings. Water Air Soil Pollution, 7, 131-136.
[25]	Yao FF (姚芳芳), Wang XK (王效科), Feng ZZ (冯兆忠), Zheng FX (郑飞翔), Feng ZW (冯宗炜), Ouyang ZY (欧阳志云) (2007). Influence of ozone and ethylenediurea (EDU) on physiological characters and foliar symptom of spinach (Spinacia oleracea L.) in open-top chambers. Ecology and Environment (生态环境), 16, 1399-1405. (in Chinese with English abstract)
[26]	Zheng QW (郑启伟), Wang XK (王效科), Feng ZZ (冯兆忠), Feng ZW (冯宗炜), Ouyang ZY (欧阳志云) (2007). Combined impact of ozone and simulated acid rain on gas exchange, growth and yield of field-grown winter wheat. Acta Scientiae Circumstantiae (环境科学学报), 27, 1542-1548. (in Chinese with English abstract)
[27]	Zheng QW (郑启伟), Wang XK (王效科), Xie JQ (谢居清), Feng ZZ (冯兆忠), Feng ZW (冯宗炜), Ni XW (倪雄伟), Ouyang ZY (欧阳志云) (2006). Effects of exogenous ascorbate acid on membrane protective system of in situ rice leaves under O₃ stress. Acta Ecologica Sinica (生态学报), 26, 1131-1137. (in Chinese with English abstract)