温度变化对油菜及其伴生杂草种苗生长和幼苗生理特性的影响

doi:10.3724/SP.J.1258.2013.00116

摘要/Abstract

摘要：

以目前大面积栽培的甘蓝型油菜(Brassica napus) ‘秦优10号’和其田间常见的8种杂草播娘蒿(Descurainia sophia)、泽漆(Euphorbia helioscopia)、野燕麦(Avena fatua)、棒头草(Polypogon fugax)、婆婆纳(Veronica didyma)、看麦娘(Alopecurus aequalis)、灰绿藜(Chenopodium glaucum)、反枝苋(Amaranthus retroflexus)为试验材料, 利用人工气候培养箱控温, 研究了10 ℃/5 ℃ (昼/夜)、18 ℃/10 ℃、25 ℃/20 ℃、35 ℃/30 ℃、40 ℃/35 ℃不同温度幅度处理对它们的发芽势、发芽率、幼苗形态及生理指标的影响, 旨在为预测气候变化环境下油菜田恶性靶标杂草的演替趋势提供参考依据。试验结果表明: 不同供试杂草对温度变化表现出明显的适应性差异, 看麦娘、灰绿藜和反枝苋的最适萌发温度偏高, 如反枝苋在高温处理(40 ℃/35 ℃)下种子的发芽率达到最高(91%); 而播娘蒿、泽漆、野燕麦、棒头草、婆婆纳种子的最适萌发温度主要集中于10-18 ℃范围内。进一步测试研究表明, 高温处理(40 ℃/35 ℃)下, 反枝苋幼苗生长旺盛, 体内丙二醛的积累也明显低于低温(10 ℃/5 ℃)处理, 可溶性糖和可溶性蛋白含量以及保护酶活性也较高, 表明反枝苋对高温胁迫有较好的适应性, 灰绿藜对高温的响应与反枝苋相似, 在高温气候年份, 要严防这2种草害加重; 而在低温处理(10 ℃/5 ℃)下, 野燕麦的种子萌发率达到100%, 其幼苗生长旺盛, 体内可溶性糖和可溶性蛋白含量及保护酶活性较高, 播娘蒿对低温胁迫的响应类似于野燕麦, 因此在寒冷年份需加强对野燕麦和播娘蒿草害的预防。供试油菜品种相对于所有供试杂草对温度变化的适应性更好, 其种子在不同温度处理下均保持较稳定的高发芽率, 且幼苗长势良好, 表明‘秦优10号’是一个对温度变化适应性很好的油菜品种。

关键词: 适应性, 油菜, 幼苗生长, 温度变化, 杂草

Abstract:

Aims Weed communities in rape fields have changed with occurrence of extreme weather events and global warming. Our objective was to investigate the effects of different regimes of temperature change on rape (Brassica napus ‘Qinyou-10’) and eight companion weeds, i.e. Descurainia sophia, Euphorbia helioscopia, Avena fatua, Polypogon fugax, Veronica didyma, Alopecurus aequalis, Chenopodium glaucum and Amaranthus retroflexus, to help with predicting the succession of companion weed communities under extreme temperatures.
Methods Seed germination, seedling growth and several physiological properties were investigated under five temperature treatments (10 °C/5 °C, 18 °C /10 °C, 25 °C /20 °C, 35 °C /30 °C and 40 °C /35 °C) (day/night) controlled by a climate-control growth chamber. Physiological properties tested included contents of malondialdehyde (MDA), soluble sugar and soluble protein, and activities of superoxide dismutase (SOD) and peroxidase (POD).
Important findings There were significant differences among the weed species in response to temperature change. The optimal temperatures for seed germination in Alopecurus aequalis, Chenopodium glaucum and Amaranthus retroflexus were high; for instance, the highest seed germination rate (91%) occurred at temperature of 40 °C /35 °C in Amaranthus retroflexus. In contrast, seed germination in Descurainia sophia, Euphorbia helioscopia, Avena fatua, Polypogon fugax and Veronica didyma required a much lower optimal temperature (i.e. 10-18 °C). At the highest treatment temperature (40 °C /35 °C), Amaranthus retroflexus seedlings grew vigorously, and their MDA contents were much lower than under low temperature treatment of 10 °C /5 °C. Moreover, the concentrations of soluble sugar and soluble protein and the activities of SOD and POD in Amaranthus retroflexus seedlings all increased with temperature, indicating the strong ability of this weed species in adapting to high temperature stress. Chenopodium glaucum had the similar responses as Amaranthus retroflexus in those properties to changes in temperature. Therefore, these two weeds should be paid more attention with the increasingly intensified global warming. Contrarily, at the lowest treatment temperature of 10 °C/5 °C, seed germination reached 100%, and the seedlings exhibited high soluble sugar and soluble protein concentrations as well as high SOD and POD activities in Avena fatua. Descurainia sophia had the similar responses as Avena fatua. Therefore, we should take additional measures to prevent malignant growth of Avena fatua and Descurainia sophia in cold years. Rape (‘Qinyou-10’) had stable and high seed germination rate, and grew well as seedlings, indicating that it better adapted to temperature changes compared to the eight companion weeds. We deduced ‘Qinyou-10’ is a rape species that can adapt well to temperature changes.

Key words: adaptability, rape, seedling growth, temperature change, weed

张凯, 慕小倩, 孙晓玉, 汪梦竹, 胡胜武. 温度变化对油菜及其伴生杂草种苗生长和幼苗生理特性的影响. 植物生态学报, 2013, 37(12): 1132-1141. DOI: 10.3724/SP.J.1258.2013.00116

ZHANG Kai, MU Xiao-Qian, SUN Xiao-Yu, WANG Meng-Zhu, HU Sheng-Wu. Effects of temperature change on seed germination, seedling growth and physiological characteristics in rape and companion weeds. Chinese Journal of Plant Ecology, 2013, 37(12): 1132-1141. DOI: 10.3724/SP.J.1258.2013.00116

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https://www.plant-ecology.com/CN/Y2013/V37/I12/1132

图/表 4

参考文献 29

[1]	Apel K, Hirt H (2004). Reactive oxygen species: metabolism, oxidative stress, and signal transduction. Annual Review of Plant Biology, 55, 373-399. DOI URL PMID
[2]	Baskin JM, Baskin CC (1977). Role of temperature in the germination ecology of three summer annual weeds. Oecologia, 30, 377-382. DOI URL PMID
[3]	Bu HY, Chen XL, Xu XL, Liu K, Jia P, Du GZ (2007). Seed mass and germination in an alpine meadow on the eastern Tsinghai-Tibet Plateau. Plant Ecology, 191, 127-149. DOI URL
[4]	Ding YH, Ren GY, Shi GY, Gong P, Zheng XH, Zhai PM, Zhang DE, Zhao ZC, Wang SW, Wang HJ, Luo Y, Chen DL, Gao XJ, Dai XS (2006). National assessment report of climate change (I): Climate change in China and its future trend. Advances in Climate Change Research, 2, 3-8. (in Chinese with English abstract)
	[ 丁一汇, 任国玉, 石广玉, 宫鹏, 郑循华, 翟盘茂, 张德二, 赵宗慈, 王绍武, 王会军, 罗勇, 陈德亮, 高学杰, 戴晓苏 (2006). 气候变化国家评估报告(I): 中国气候变化的历史和未来趋势. 气候变化研究进展, 2, 3-8.]
[5]	Feng ZZ, Zhou HY, Feng ZW, Wang XK (2005). Effect of triadimefon on physiological characters in leaves of cucumber seedlings under high temperature. Acta Botanica Boreali-Occidentalia Sinica, 25, 170-173. (in Chinese with English abstract)
	[ 冯兆忠, 周华英, 冯宗炜, 王效科 (2005). 高温胁迫下三唑酮对黄瓜幼苗某些生理性质的影响. 西北植物学报, 25, 170-173.]
[6]	Gao JF (2006). Plant Physiology Experimental Guidance. Higher Education Press, Beijing. (in Chinese)
	[ 高俊凤 (2006). 植物生理学实验指导. 高等教育出版社, 北京.]
[7]	Guan B, Zhou DW, Tian Y, Yang JY, Xiao MX (2010). Effect of salinity, alkalinity and temperature on germination of Medicago ruthenica seeds. Chinese Journal of Grassland, 32(1), 58-63. (in Chinese with English abstract)
	[ 管博, 周道玮, 田雨, 杨季云, 肖模昕 (2010). 盐碱及变温条件对花苜蓿种子发芽的影响. 中国草地学报, 32(1), 58-63.]
[8]	Hou FL (2004). Tutorial of Plant Physiology Experiment. Science Press, Beijing. (in Chinese)
	[ 侯福林 (2004). 植物生理学实验教程. 科学出版社, 北京.]
[9]	Khan MA, Guhar S (2003). Germination responses of Sporobolus ioclados: a saline desert grass. Journal of Arid Environments, 53, 387-394. DOI URL
[10]	Lin JX, Li XY, Tang JH, Wang JF, Li ZL, Zhang ZJ, Mu CS (2011). Effects of temperature, salinity, alkalinity and their interactions on seed germination of wheat. Crops, (6), 113-116. (in Chinese with English abstract)
	[ 蔺吉祥, 李晓宇, 唐佳红, 王俊锋, 李卓琳, 张兆军, 穆春生 (2011). 温度与盐、碱胁迫交互作用对小麦种子萌发的影响. 作物杂志, (6), 113-116.]
[11]	Liu P, Li MJ (2007). Plant Physiology Experimental Techniques. Science Press, Beijing. (in Chinese)
	[ 刘萍, 李明军 (2007). 植物生理学实验技术. 科学出版社, 北京.]
[12]	Liu W, Liu K, Zhang CH, Du GZ (2011). Effect of accumulated temperature on seed germination—a case study of 12 Compositae species on the eastern Qinghai-Tibet Plateau of China. Chinese Journal of Plant Ecology, 35, 751-758. (in Chinese with English abstract) DOI URL
	[ 刘文, 刘坤, 张春辉, 杜国祯 (2011). 种子萌发的积温效应——以青藏高原东缘的12种菊科植物为例. 植物生态学报, 35, 751-758.]
[13]	Luo H (2010). Situation and development prospect of rape in Shaanxi Province. Shanxi Journal of Agricultural Sciences, (2), 114-116. (in Chinese with English abstract)
	[ 罗恒 (2010). 陕西省油菜生产现状与发展前景. 陕西农业科学, (2), 114-116.]
[14]	Ma HY, Liang ZW, Kong XJ, Yan C, Chen Y (2008). Effects of salinity, temperature and their interaction on the germination percentage and seedling growth of Leymus chinensis (Trin.) Tzve1. (Poaceae). Acta Ecologica Sinica, 28, 4710-4717. (in Chinese with English abstract)
	[ 马红媛, 梁正伟, 孔祥军, 闫超, 陈渊 (2008). 盐分、温度及其互作对羊草种子发芽率和幼苗生长的影响. 生态学报, 28, 4710-4717.]
[15]	Mohsenzadeh S, Malboobi MA, Razavi K, Farrahi-Aschtiani S (2006). Physiological and molecular responses of Aeluropus lagopoides (Poaceae) to water deficit. Environmental and Experimental Botany, 56, 314-322. DOI URL
[16]	Norden N, Daws MI, Antoine C, Gonzalez MA, Garwood NC, Chave J (2009). The relationship between seed mass and mean time to germination for 1037 tree species across five tropical forests. Functional Ecology, 23, 203-210. DOI URL
[17]	Qian LH, Ruan SL, Dai DL, Xin Y (2010). Effect of temperature on SOD, MDA of Radix Tetrastigmae. Journal of Zhejiang Agricultural Sciences, (5), 972-974. (in Chinese)
	[ 钱丽华, 阮松林, 戴丹丽, 忻雅 (2010). 温度对三叶青组培苗SOD、MDA等指标的影响. 浙江农业科学, (5), 972-974.]
[18]	Robers EH (1988). Temperature and seed germination. Symposia of the Society for Experimental Biology, 42, 109-132. URL PMID
[19]	Tlig T, Gorai M, Neffati M (2008). Germination responses of Diplotaxis harra to temperature and salinity. Flora-Morphology, Distribution, Functional Ecology of Plants, 203, 421-428.
[20]	Wang RY, Li RZ, Sun ZY, Ren YS, Yue WB (2006). Anti freezing proteins and plant responses to low temperature stress. Chinese Journal of Applied Ecology, 17, 551-556 (in Chinese with English abstract)
	[ 王瑞云, 李润植, 孙振元, 任有蛇, 岳文斌 (2006). 抗冻蛋白与植物低温胁迫反应. 应用生态学报, 17, 551-556.]
[21]	Yang D, Zhang H, Chen LP, Ma DW (2007). Effect of temperature stress on propanedial and soluble sugar contents in ten Compositae weeds. Journal of Sichuan Normal University (Natural Science), 30, 391-394. (in Chinese with English abstract)
	[ 杨东, 张红, 陈丽萍, 马丹炜 (2007). 温度胁迫对10种菊科杂草丙二醛和可溶性糖的影响. 四川师范大学学报(自然科学版), 30, 391-394.]
[22]	Yang HG, Chen HJ (2009). Effect of high temperature stress on morphological and physiological characteristics in Phalaenopsis seedlings. Chinese Agricultural Science Bulletin, 25, 123-127. (in Chinese with English abstract)
	[ 杨华庚, 陈慧娟 (2009). 高温胁迫对蝴蝶兰幼苗叶片形态和生理特性的影响. 中国农学通报, 25, 123-127.]
[23]	Yang SS, Gao JF (2001). Influence of active of oxygen and free radicals on plant senescence. Acta Botanica Boreali-Occidentalia Sinica, 21, 215-220. (in Chinese with English abstract)
	[ 杨淑缜, 高俊凤 (2001). 活性氧、自由基与植物的衰老. 西北植物学报, 21, 215-220.]
[24]	Zeng GW, Jiang DA (2000). Plant Physiology. China Agricultural Science and Technology Press, Beijing. (in Chinese)
	[ 曾广文, 蒋德安 (2000). 植物生理学. 中国农业科技出版社, 北京.]
[25]	Zhang FY, Lei G, Zhang P, Sun CY (2012). Effects of salicylic acid on the germination of velvetbean seeds and phyological characteristics of velvetbean seedlings under cold stress. Journal of Northwest A & F University (Natural Science Edition), 40(4), 205-209, 216.
	[ 张凤银, 雷刚, 张萍, 孙彩云 (2012). 水杨酸对低温胁迫下藜豆种子萌发和幼苗生理特性的影响. 西北农林科技大学学报(自然科学版), 40(4), 205-209, 216.]
[26]	Zhang HX (2008). Research on Seed Germination Ecology. PhD dissertation, Northeast Normal University, Changchun. (in Chinese with English abstract)
	[ 张红香 (2008). 种子发芽生态研究. 博士学位论文, 东北师范大学, 长春.]
[27]	Zhang Y, Mu XQ (2008). Allelopathic effects of Amaranthus retroflexus L. and its risk assessment. Acta Botanica Boreali-Occidentalia Sinica, 28, 771-776. (in Chinese with English abstract)
	[ 张燕, 慕小倩 (2008). 外来杂草反枝苋对农作物的化感作用及其风险评价. 西北植物学报, 28, 771-776.]
[28]	Zhang YP, Yang SJ, Chen FX, Chen YY (2011). Effects of salinity, temperature and their interaction on seed germination performance and seedling growth of melon. Seed, 30(10), 15-18, 24. (in Chinese with English abstract)
	[ 张永平, 杨少军, 陈绯翔, 陈幼源 (2011). 盐分、温度及其互作对甜瓜种子萌发及幼苗生长的影响. 种子, 30(10), 15-18, 24.]
[29]	Zhu WD, Wei SH, Liu X, Zhang CX, Zhang HJ, Zhang J, Yu DZ (2007). Occurrence rule and chemical control of weed community in rape field. Hubei Agricultural Sciences, 46, 936-938. (in Chinese with English abstract)
	[ 朱文达, 魏守辉, 刘学, 张朝贤, 张宏军, 张佳, 喻大昭 (2007). 油菜田杂草发生规律及化学防除技术. 湖北农业科学, 46, 936-938.]

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	发芽势 Germination energy (%)					发芽率 Germination rate (%)
	T1	T2	T3	T4	T5	T1	T2	T3	T4	T5
播娘蒿 Descurainia sophia	42^b	80^a	11^c	10^c	10^c	52^b	92^a	20^c	16^d	14^d
泽漆 Euphorbia helioscopia	20^b	31^a	36^a	20^b	16^c	35^a	40^a	40^a	35^a	22^b
野燕麦 Avena fatua	80^a	90^a	90^a	10^b	5^c	100^a	100^a	100^a	20^b	12^c
棒头草 Polypogon fugax	16^e	70^a	62^b	40^d	56^c	32^c	87^b	90^a	91^a	90^a
婆婆纳 Veronica didyma	32^c	87^a	60^b	65^b	60^b	62^d	100^a	95^b	90^c	91^c
看麦娘 Alopecurus aequalis	2^c	3^b	6^a	3^b	6^a	3^d	21^c	25^b	26^b	35^a
灰绿藜 Chenopodium glaucum	5^c	4^c	3^c	10^b	19^a	6^e	10^d	12^c	15^b	26^a
反枝苋 Amaranthus retroflexus	2^e	4^d	16^c	60^b	85^a	3^e	7^d	20^c	85^b	91^a
油菜 Brassica napus	81^c	90^b	96^a	84^c	62^d	88^b	98^a	100^a	98^a	100^a

	发芽势 Germination energy (%)					发芽率 Germination rate (%)
	T1	T2	T3	T4	T5	T1	T2	T3	T4	T5
播娘蒿 Descurainia sophia	42^b	80^a	11^c	10^c	10^c	52^b	92^a	20^c	16^d	14^d
泽漆 Euphorbia helioscopia	20^b	31^a	36^a	20^b	16^c	35^a	40^a	40^a	35^a	22^b
野燕麦 Avena fatua	80^a	90^a	90^a	10^b	5^c	100^a	100^a	100^a	20^b	12^c
棒头草 Polypogon fugax	16^e	70^a	62^b	40^d	56^c	32^c	87^b	90^a	91^a	90^a
婆婆纳 Veronica didyma	32^c	87^a	60^b	65^b	60^b	62^d	100^a	95^b	90^c	91^c
看麦娘 Alopecurus aequalis	2^c	3^b	6^a	3^b	6^a	3^d	21^c	25^b	26^b	35^a
灰绿藜 Chenopodium glaucum	5^c	4^c	3^c	10^b	19^a	6^e	10^d	12^c	15^b	26^a
反枝苋 Amaranthus retroflexus	2^e	4^d	16^c	60^b	85^a	3^e	7^d	20^c	85^b	91^a
油菜 Brassica napus	81^c	90^b	96^a	84^c	62^d	88^b	98^a	100^a	98^a	100^a

		播娘蒿 Descurainia sophia	泽漆 Euphorbia helioscopia	野燕麦 Avena fatua	棒头草 Polypogon fugax	婆婆纳 Veronica didyma	看麦娘 Alopecurus aequalis	灰绿藜 Chenopodium glaucum	反枝苋 Amaranthus retroflexus
丙二醛 Malonaldehyde (nmol·g^-1 FW)	T1	4.98 ± 0.64^ab	3.41 ± 0.14^b	1.31 ± 0.22^a	5.29 ± 0.95^a	5.21 ± 1.02^b	46.31 ± 1.80^a	1.75 ± 0.09^a	10.18 ± 0.63^a
	T2	2.80 ± 0.09^c	2.42 ± 0.08^b	0.97 ± 0.45^b	2.35 ± 0.24^b	4.13 ± 1.42^b	10.61 ± 0.80^bc	1.45 ± 0.12^ab	8.44 ± 0.52^b
	T3	3.73 ± 0.15^bc	0.36 ± 0.01^c	0.27 ± 0.22^c	1.63 ± 0.05^b	5.12 ± 0.31^b	8.80 ± 1.45^c	1.27 ± 0.02^b	5.59 ± 0.01^c
	T4	4.05 ± 0.96^abc	2.36 ± 0.25^b	0.30 ± 0.49^c	1.75 ± 0.22^b	6.58 ± 0.31^b	15.35 ± 0.43^bc	0.73 ± 0.05^c	4.69 ± 0.13^c
	T5	5.56 ± 0.12^a	6.10 ± 0.68^a	0.35 ± 0.02^c	4.62 ± 0.07^a	9.53 ± 1.47^a	23.80 ± 0.15^b	1.51 ± 0.20^ab	4.97 ± 0.14^c
可溶性糖 Soluble sugar (mg·g^-1 FW)	T1	55.67 ± 0.31^a	152.89 ± 0.50^b	70.47 ± 0.22^a	80.70 ± 0.09^a	299.47 ± 0.02^ab	511.55 ± 0.24^d	42.35 ± 0.02^b	82.93 ± 0.39^a
	T2	50.05 ± 0.37^ab	117.56 ± 0.42^c	57.23 ± 0.45^b	70.30 ± 0.44^b	201.60 ± 1.14^c	621.08 ± 3.12^b	39.37 ± 0.42^a	68.71 ± 0.32^b
	T3	39.57 ± 0.61^b	60.42 ± 0.03^d	29.87 ± 0.05^c	36.76 ± 0.01^d	223.91 ± 0.05^c	562.93 ± 0.20^c	10.20 ± 0.05^b	53.94 ± 0.29^c
	T4	42.91 ± 0.20^ab	105.60 ± 0.59^c	33.45 ± 0.02^c	61.27 ± 0.24^e	253.65 ± 3.71^bc	737.33 ± 2.06^a	34.73 ± 0.31^a	72.30 ± 0.22^b
	T5	54.48 ± 1.16^ab	241.21 ± 1.53^a	34.50 ± 0.02^c	102.50 ± 0.05^c	329.000 ± 0.03^a	398.82 ± 0.44^e	10.21 ± 0.02^b	76.72 ± 0.24^ab
可溶性蛋白 Soluble protein (mg·g^-1FW)	T1	9.66 ± 0.14^e	12.96 ± 0.14^c	15.62 ± 0.21^b	20.10 ± 0.33^b	4.56 ± 0.15^b	33.42 ± 0.87^b	11.00 ± 0.03^c	21.01 ± 0.75^b
	T2	16.64 ± 0.11^b	13.92 ± 0.12^b	16.64 ± 0.19^a	17.37 ± 0.08^c	4.91 ± 0.07^b	9.99 ± 0.20^d	17.16 ± 0.13^a	19.75 ± 0.04^c
	T3	11.63 ± 0.07^d	10.58 ± 0.19^d	9.60 ± 0.14^e	10.60 ± 0.20^e	3.35 ± 0.26^c	9.69 ± 0.42^d	6.73 ± 0.10^d	16.78 ± 0.23^d
	T4	20.40 ± 0.22^a	16.16 ± 0.08^a	10.92 ± 0.16^d	11.62 ± 0.11^d	3.28 ± 0.38^c	29.52 ± 0.55^c	16.63 ± 0.15^b	21.87 ± 0.14^b
	T5	12.18 ± 0.07^c	9.63 ± 0.20^e	14.25 ± 0.08^c	28.00 ± 0.11^a	13.97 ± 0.14^a	50.76 ± 0.73^a	11.17 ± 0.05^c	25.68 ± 0.25^a
超氧化物歧化酶 Superoxide dismutase U·g^-1 FW·min^-1	T1	258.70 ± 3.39^b	323.77 ± 6.89^b	130.75 ± 2.33^a	121.15 ± 6.48^a	421.88 ± 1.04^b	317.95 ± 6.22^b	456.37 ± 1.08^b	199.20 ± 10.14^b
	T2	126.04 ± 1.39^d	184.54 ± 0.75^c	65.62 ± 1.06^c	92.58 ± 1.97^bc	94.76 ± 0.52^e	119.82 ± 9.50^d	206.69 ± 8.65^d	59.31 ± 3.41^c
	T3	50.47 ± 1.94^e	51.33 ± 11.12^d	29.65 ± 4.11^d	68.51 ± 2.43^d	131.03 ± 6.13^d	104.27 ± 3.15^d	63.08 ± 1.84^e	78.84 ± 3.97^c
	T4	228.43 ± 6.22^c	171.35 ± 1.50^c	70.50 ± 10.35^c	80.02 ± 10.69^cd	220.27 ± 11.01^c	143.24 ± 2.45^c	374.39 ± 3.17^c	199.87 ± 3.68^b
	T5	337.23 ± 1.91^a	352.63 ± 1.92^a	91.20 ± 2.62^b	101.77 ± 2.35^b	463.63 ± 8.03^a	500.75 ± 6.98^a	472.60 ± 1.10^a	551.15 ± 6.23^a
过氧化物酶 Peroxidase U·g^-1 FW·min^-1	T1	348.37 ± 5.52^c	451.48 ± 1.52^c	1203.67 ± 18.18^a	450.99 ± 4.52^b	487.79 ± 2.56^a	63.53 ± 6.64^c	147.79 ± 0.54^b	182.59 ± 6.75^b
	T2	709.55 ± 2.76^b	346.97 ± 4.33^d	979.87 ± 2.61^b	341.28 ± 8.36^c	70.59 ± 1.81^d	55.88 ± 1.86^c	70.76 ± 1.48^e	219.972 ± 6.52^ab
	T3	356.38 ± 1.40^c	327.98 ± 0.32^e	633.91 ± 13.43^e	253.00 ± 9.35^c	106.68 ± 1.88^c	57.03 ± 4.47^c	81.17 ± 0.28^d	173.28 ± 1.90^b
	T4	851.68 ± 1.82^a	745.16 ± 9.70^b	748.65 ± 16.32^d	723.16 ± 2.12^a	293.68 ± 6.08^b	86.47 ± 4.99^b	161.50 ± 2.40^a	192.78 ± 1.79^b
	T5	257.59 ± 2.21^d	831.68 ± 2.87^a	805.43 ± 6.62^d	775.99 ± 5.13^a	514.87 ± 2.05^a	118.44 ± 1.29^a	118.28 ± 1.61^c	266.70 ± 6.90^a

		播娘蒿 Descurainia sophia	泽漆 Euphorbia helioscopia	野燕麦 Avena fatua	棒头草 Polypogon fugax	婆婆纳 Veronica didyma	看麦娘 Alopecurus aequalis	灰绿藜 Chenopodium glaucum	反枝苋 Amaranthus retroflexus
丙二醛 Malonaldehyde (nmol·g^-1 FW)	T1	4.98 ± 0.64^ab	3.41 ± 0.14^b	1.31 ± 0.22^a	5.29 ± 0.95^a	5.21 ± 1.02^b	46.31 ± 1.80^a	1.75 ± 0.09^a	10.18 ± 0.63^a
	T2	2.80 ± 0.09^c	2.42 ± 0.08^b	0.97 ± 0.45^b	2.35 ± 0.24^b	4.13 ± 1.42^b	10.61 ± 0.80^bc	1.45 ± 0.12^ab	8.44 ± 0.52^b
	T3	3.73 ± 0.15^bc	0.36 ± 0.01^c	0.27 ± 0.22^c	1.63 ± 0.05^b	5.12 ± 0.31^b	8.80 ± 1.45^c	1.27 ± 0.02^b	5.59 ± 0.01^c
	T4	4.05 ± 0.96^abc	2.36 ± 0.25^b	0.30 ± 0.49^c	1.75 ± 0.22^b	6.58 ± 0.31^b	15.35 ± 0.43^bc	0.73 ± 0.05^c	4.69 ± 0.13^c
	T5	5.56 ± 0.12^a	6.10 ± 0.68^a	0.35 ± 0.02^c	4.62 ± 0.07^a	9.53 ± 1.47^a	23.80 ± 0.15^b	1.51 ± 0.20^ab	4.97 ± 0.14^c
可溶性糖 Soluble sugar (mg·g^-1 FW)	T1	55.67 ± 0.31^a	152.89 ± 0.50^b	70.47 ± 0.22^a	80.70 ± 0.09^a	299.47 ± 0.02^ab	511.55 ± 0.24^d	42.35 ± 0.02^b	82.93 ± 0.39^a
	T2	50.05 ± 0.37^ab	117.56 ± 0.42^c	57.23 ± 0.45^b	70.30 ± 0.44^b	201.60 ± 1.14^c	621.08 ± 3.12^b	39.37 ± 0.42^a	68.71 ± 0.32^b
	T3	39.57 ± 0.61^b	60.42 ± 0.03^d	29.87 ± 0.05^c	36.76 ± 0.01^d	223.91 ± 0.05^c	562.93 ± 0.20^c	10.20 ± 0.05^b	53.94 ± 0.29^c
	T4	42.91 ± 0.20^ab	105.60 ± 0.59^c	33.45 ± 0.02^c	61.27 ± 0.24^e	253.65 ± 3.71^bc	737.33 ± 2.06^a	34.73 ± 0.31^a	72.30 ± 0.22^b
	T5	54.48 ± 1.16^ab	241.21 ± 1.53^a	34.50 ± 0.02^c	102.50 ± 0.05^c	329.000 ± 0.03^a	398.82 ± 0.44^e	10.21 ± 0.02^b	76.72 ± 0.24^ab
可溶性蛋白 Soluble protein (mg·g^-1FW)	T1	9.66 ± 0.14^e	12.96 ± 0.14^c	15.62 ± 0.21^b	20.10 ± 0.33^b	4.56 ± 0.15^b	33.42 ± 0.87^b	11.00 ± 0.03^c	21.01 ± 0.75^b
	T2	16.64 ± 0.11^b	13.92 ± 0.12^b	16.64 ± 0.19^a	17.37 ± 0.08^c	4.91 ± 0.07^b	9.99 ± 0.20^d	17.16 ± 0.13^a	19.75 ± 0.04^c
	T3	11.63 ± 0.07^d	10.58 ± 0.19^d	9.60 ± 0.14^e	10.60 ± 0.20^e	3.35 ± 0.26^c	9.69 ± 0.42^d	6.73 ± 0.10^d	16.78 ± 0.23^d
	T4	20.40 ± 0.22^a	16.16 ± 0.08^a	10.92 ± 0.16^d	11.62 ± 0.11^d	3.28 ± 0.38^c	29.52 ± 0.55^c	16.63 ± 0.15^b	21.87 ± 0.14^b
	T5	12.18 ± 0.07^c	9.63 ± 0.20^e	14.25 ± 0.08^c	28.00 ± 0.11^a	13.97 ± 0.14^a	50.76 ± 0.73^a	11.17 ± 0.05^c	25.68 ± 0.25^a
超氧化物歧化酶 Superoxide dismutase U·g^-1 FW·min^-1	T1	258.70 ± 3.39^b	323.77 ± 6.89^b	130.75 ± 2.33^a	121.15 ± 6.48^a	421.88 ± 1.04^b	317.95 ± 6.22^b	456.37 ± 1.08^b	199.20 ± 10.14^b
	T2	126.04 ± 1.39^d	184.54 ± 0.75^c	65.62 ± 1.06^c	92.58 ± 1.97^bc	94.76 ± 0.52^e	119.82 ± 9.50^d	206.69 ± 8.65^d	59.31 ± 3.41^c
	T3	50.47 ± 1.94^e	51.33 ± 11.12^d	29.65 ± 4.11^d	68.51 ± 2.43^d	131.03 ± 6.13^d	104.27 ± 3.15^d	63.08 ± 1.84^e	78.84 ± 3.97^c
	T4	228.43 ± 6.22^c	171.35 ± 1.50^c	70.50 ± 10.35^c	80.02 ± 10.69^cd	220.27 ± 11.01^c	143.24 ± 2.45^c	374.39 ± 3.17^c	199.87 ± 3.68^b
	T5	337.23 ± 1.91^a	352.63 ± 1.92^a	91.20 ± 2.62^b	101.77 ± 2.35^b	463.63 ± 8.03^a	500.75 ± 6.98^a	472.60 ± 1.10^a	551.15 ± 6.23^a
过氧化物酶 Peroxidase U·g^-1 FW·min^-1	T1	348.37 ± 5.52^c	451.48 ± 1.52^c	1203.67 ± 18.18^a	450.99 ± 4.52^b	487.79 ± 2.56^a	63.53 ± 6.64^c	147.79 ± 0.54^b	182.59 ± 6.75^b
	T2	709.55 ± 2.76^b	346.97 ± 4.33^d	979.87 ± 2.61^b	341.28 ± 8.36^c	70.59 ± 1.81^d	55.88 ± 1.86^c	70.76 ± 1.48^e	219.972 ± 6.52^ab
	T3	356.38 ± 1.40^c	327.98 ± 0.32^e	633.91 ± 13.43^e	253.00 ± 9.35^c	106.68 ± 1.88^c	57.03 ± 4.47^c	81.17 ± 0.28^d	173.28 ± 1.90^b
	T4	851.68 ± 1.82^a	745.16 ± 9.70^b	748.65 ± 16.32^d	723.16 ± 2.12^a	293.68 ± 6.08^b	86.47 ± 4.99^b	161.50 ± 2.40^a	192.78 ± 1.79^b
	T5	257.59 ± 2.21^d	831.68 ± 2.87^a	805.43 ± 6.62^d	775.99 ± 5.13^a	514.87 ± 2.05^a	118.44 ± 1.29^a	118.28 ± 1.61^c	266.70 ± 6.90^a