砧穗互作对菊花嫁接苗耐盐性的影响

doi:10.3864/j.issn.0578-1752.2021.03.016

摘要/Abstract

摘要：

【目的】揭示黄蒿嫁接菊花对菊花耐盐性的影响,明确砧穗互作影响菊花耐盐性的机理。【方法】测定120 mmol·L^-1NaCl胁迫下的‘钟山嫣红’的扦插苗（自根苗）、扦插苗的自根嫁接苗（自接苗）、黄蒿砧木嫁接菊花（异根嫁接苗）的生理指标和叶片、茎段的Na⁺、K⁺离子含量。【结果】NaCl胁迫下,异根嫁接苗叶片受害率低于‘钟山嫣红’的自根苗和自接苗。异根嫁接苗的叶片相对电导率、丙二醛含量、脯氨酸含量均低于自接苗和自根苗,叶绿素含量、超氧化物歧化酶（SOD）、过氧化物酶（POD）含量高于自接苗和自根苗。异根嫁接苗中部与下部叶的Na⁺含量、Na⁺/K⁺比值均最低,其次是自接苗,最高是自根苗。自根嫁接与异根嫁接苗上部叶的Na⁺含量差异不显著;异根嫁接苗接口以上茎段的Na⁺含量显著低于自接苗和自根苗,而自接苗和自根苗上部茎段的Na⁺无显著差异,接口以下的黄蒿茎段Na⁺浓度和Na⁺/K⁺是同为砧木的菊花茎段中的两倍。【结论】菊花自接苗运输到中部、下部叶片的离子显著少于自根苗,以黄蒿作为砧木嫁接菊花后,砧木对Na⁺较高的富集能力降低了Na⁺在嫁接接口上部的积累,同时,菊花异根嫁接苗表现出更强的光合性能和抗氧化酶活性。因此,以黄蒿作砧木的‘钟山嫣红’异根嫁接苗耐盐性提高,是砧穗愈合导致向上运输到叶片的离子减少和黄蒿砧木滞留更多Na⁺的共同作用。

关键词: 嫁接, 菊花, 黄蒿, 生理指标, Na⁺, K⁺, 耐盐性

Abstract:

【Objective】The objective of this study is to investigate the effects of Artemisia annua grafted with chrysanthemum on the salt tolerance and its interaction between the rootstocks and scions. 【Method】The physiological indexes, including the content of Na⁺ and K⁺ in the leaves and stems of chrysanthemum cutting seedlings Zhongshan Yanhong (self-rooted seedlings), self-root grafted cutting seedlings (self-grafted seedlings) and Artemisia annua rootstock grafted cutting seedlings (hetero-root grafted seedlings), were measured under 120 mmol·L^-1NaCl stress. 【Result】Under NaCl stress, the damage symptom rate of hetero-root grafted seedling leaves was lower than that of self-rooted and self-grafted seedlings. The relative content of conductivity, malondialdehyde, and proline of hetero-root grafted seedlings were lower than those of self-rooted and self-grafted seedlings, and the content of chlorophyll, superoxide dismutase (SOD), and peroxidase (POD) were higher than self-rooted and self-grafted seedlings. The content of Na⁺ and Na⁺/K⁺ ratios of the middle and lower leaves of the hetero-root grafted seedlings showed the lowest, followed by the self-grafted seedlings, and the highest index was recorded in the self-rooted seedlings. There was no significant difference in Na⁺ content between the upper leaves of self-grafted seedlings and hetero-root grafted seedlings; the Na⁺ content of the stems above the interface of hetero-root grafted seedlings was significantly lower than that of self-grafted and self-rooted seedlings, while there was no significant difference in the Na⁺ content of the upper stems of self-grafted and self-rooted seedlings; the Na⁺ concentration and Na⁺/K⁺ of Artemisia annua stem segments below the interface were twice fold than that of the stem segments of chrysanthemum.【Conclusion】The ion transported from self-grafted seedlings of chrysanthemum to the middle and lower leaves was significantly less than that of self-rooted seedlings. The results indicated that after grafting chrysanthemum with Artemisia annua as a rootstock, the higher Na⁺ enrichment capacity of rootstock reduced the accumulation of Na⁺ in the upper part of the grafting interface, while hetero-root grafted seedlings showed stronger photosynthetic performance and antioxidant enzyme activity. Therefore, the improvement of salt tolerance of hetero-root grafted seedlings with Artemisia annua as rootstock was the combined effect of rootstock healing leading to the reduction of ions transported upward to the leaves and the accumulation of Artemisia annua rootstock with more Na⁺ on the rootstock.

Key words: grafting, chrysanthemum, Artemisia annua, physiological index, Na⁺, K⁺, salt tolerance

孟蕊,刘晔,赵爽,房伟民,蒋甲福,陈素梅,陈发棣,管志勇. 砧穗互作对菊花嫁接苗耐盐性的影响[J]. 中国农业科学, 2021, 54(3): 629-642.

MENG Rui,LIU Ye,ZHAO Shuang,FANG WeiMin,JIANG JiaFu,CHEN SuMei,CHEN FaDi,GUAN ZhiYong. Effects of Rootstock and Scion Interaction on Salt Tolerance of Grafted Chrysanthemum Seedlings[J]. Scientia Agricultura Sinica, 2021, 54(3): 629-642.

0
/ / 推荐

导出引用管理器 EndNote|Reference Manager|ProCite|BibTeX|RefWorks

链接本文: https://www.chinaagrisci.com/CN/10.3864/j.issn.0578-1752.2021.03.016

https://www.chinaagrisci.com/CN/Y2021/V54/I3/629

图/表 11

表1

图1

图2

图3

图4

表2

图5

图6

图7

图8

图9

参考文献 51

[1]	郝洪波. 地被小菊的特点与园林应用. 南方农业: 园林花卉版, 2007,1(6):38-40.
	HAO H B. Characteristics and ground application of ground cover chrysanthemum. South China Agriculture, 2007,1(6):38-40. (in Chinese)
[2]	房伟民. 嫁接对菊花抗逆性影响及切花菊栽培技术研究[D]. 南京: 南京农业大学, 2009.
	FANG W M. Effect on abiotic stress tolerance of grafting in chrysanthemum and cultivated techniques for cutting chrysanthemum[D]. Nanjing: Nanjing Agricultural University, 2009. (in Chinese)
[3]	ZHANG J L, SHI H Z. Physiological and molecular mechanisms of plant salt tolerance. Photosynthesis Research, 2013,115(1):1-22. doi: 10.1007/s11120-013-9813-6 pmid: 23539361
[4]	MUNNS R, TESTER M. Mechanisms of salinity tolerance. Annual Review of Plant Biology, 2008,59(1):651-681.
[5]	ZHU J K. Plant salt tolerance. Trends in Plant Science, 2001,6(2):66-71.
[6]	KRONZUCHER H J, BRITTO D T. Sodium transport in plants: Acritical review. New Phytologist, 2011,189(1):54-81.
[7]	SZABADOS L, SAVOURE A. Proline: A multifunctional amino acid. Trends in Plant Science, 2010,15(2):89-97. doi: 10.1016/j.tplants.2009.11.009 pmid: 20036181
[8]	朱士农, 郭世荣. 嫁接对盐胁迫下西瓜植株体内Na⁺和K⁺含量及其分布的影响 . 园艺学报, 2009,36(6):814-820.
	ZHU S N, GUO S R. Effects of grafting on ⁺, Na⁺ contents and distribution of watermelon (Citrullus vulgaris Schrad) seedlings under NaCl stress . Acta Horiculture Sinica, 2009,36(6):814-820. (in Chinese)
[9]	EETAN MARIA T, MARTINEZ-RODRIGUEZ M M, PEREZ-ALFOCEA F, FLOWERS T J, BOLARIN M C . Grafting raises the salt tolerance of tomato through limiting the transport of sodium and chloride to the shoot. Journal of Experimental Botany, 2005,56(412):703-712. doi: 10.1093/jxb/eri027 pmid: 15557292
[10]	雷波. 耐盐砧木通过调节Na⁺吸收和运转提高黄瓜耐盐性的机制研究 [D]. 武汉: 华中农业大学, 2013.
	LEI B. Mechanism of salt tolerant rootstock raise the salt tolerance of cucumber through regulating the uptake transportation of Na⁺ [D]. Wuhan: Huazhong Agricultural University, 2013. (in Chinese)
[11]	MARTINEZ-RODRIGUEZ M M, ESTA M T, MOYANO E, GARCIA-ABELLANT J O, FLORES F B, CAMPOS J F, AL- AZZAWI M J, FLOWERS T J, BOLARIN M C. The effectiveness of grafting to improve salt tolerance in tomato when an ‘excluder’ genotype is used as scion. Environmental & Experimental Botany, 2008,63(1):392-401.
[12]	杨秀玲. 黄瓜嫁接幼苗耐盐光合特性和WSC代谢生理调控研究[D]. 兰州: 甘肃农业大学, 2015.
	YANG X L. Physiological regulation of salt tolerance on photosynthesis and WSC metabolism in grafted cucumber seedlings[D]. Lanzhou: Gansu Agricultural University, 2015. (in Chinese)
[13]	PENELLA C, LANDI M, GUIDI L, NEBAUER S G, SAN BAUTISTA A, REMIRINI D, NALI C, LOPEZ-GALARZA S, CALATAYUD A. 盐胁迫下耐盐砧木通过保持光合能力和库强提高嫁接辣椒的产量. 刘越, 孔秋生, 译. 辣椒杂志, 2016,14(2):38-50.
	PENELLA C, LANDI M, GUIDI L, NEBAUER S G, SAN BAUTISTA A, REMIRINI D, NALI C, LOPEZ-GALARZA S, CALATAYUD A. Salt-tolerant rootstocks increase the yield of grafted peppers by maintaining photosynthetic capacity and storage strength under salt stress. LIU Y, KONG Q S, Trans. Journal of China Capsicum, 2016,14(2):38-50. (in Chinese)
[14]	高俊杰, 张琳, 秦爱国, 于贤昌. 氯化钠胁迫下嫁接黄瓜叶片SOD和CAT mRNA基因表达及其活性. 应用生态学报, 2008,19(8):1754-1758.
	GAO J J, ZHANG L, QIN A G, YU X C. Relative expression of SOD and CAT mRNA and activities of SOD and CAT in grafted cucumber leaves under NaCl stress. Chinese Journal of Applied Ecology, 2008,19(8):1754-1758. (in Chinese)
[15]	施先锋. 南瓜砧木嫁接提高西瓜耐冷性的生理机制及蛋白质组学研究[D]. 武汉: 华中农业大学, 2019.
	SHI X F. Physiological mechanism and proteomics research of improved cold tolerance of watermelon seedlings by grafting onto pumpkin rootstock[D]. Wuhan: Huazhong Agricultural University, 2019. (in Chinese)
[16]	SHAHID M A, BALAL R M, KHAN N, SIMON-GRAO S, ALFOSEA-SIMON M, CAMARA-ZAPATA J M, MATTSON N S, GARCIA-SANCHEZ F. Rootstocks influence the salt tolerance of Kinnow mandarin trees by altering the antioxidant defense system, osmolyte concentration, and toxic ion accumulation. Scientia Horticulturae, 2019,250:1-11.
[17]	林有润. 中国蒿属志—中国蒿属植物的系统分类、分布和主要经济用途. 植物研究, 1988,8(4):1-61
	LIN Y R. The Chinese Artemisia Linn. -The classification, distribution and application of Artemisia Linn In China. Bulletin of Botanical Research, 1988,8(4):1-61. (in Chinese)
[18]	CHEN Y, SUN X Z, ZHENG C S, ZHANG S, YANG J H. Grafting onto Artemisia annua improves drought tolerance in chrysanthemum by enhancing photosynthetic capacity. Horticultural Plant Journal, 2018,4(3):117-125.
[19]	ZHANG X Y, SUN X Z, ZHANG S, YANG J H, LIU F F, FAN J. Comprehensive transcriptome analysis of grafting onto Artemisia scoparia W. to affect the aphid resistance of chrysanthemum (Chrysanthemum morifolium T.). BMC Genomics, 2019,20(1):776. doi: 10.1186/s12864-019-6158-3 pmid: 31653200
[20]	陈思思. 嫁接缓解菊花连作障碍机制初探[D]. 南京: 南京农业大学, 2012.
	CHEN S S. Effects of grafting on the relieve of continuous cropping obstacles of Chrysanthemum [D]. Nanjing: Nanjing Agricultural University, 2012.
[21]	管志勇, 陈素梅, 王艳艳, 陈发棣. 菊花近缘种属植物耐盐筛选浓度的确定及耐盐性比较. 生态学杂志, 2010,29(3):467-472.
	GUAN Z Y, CHEN S M, WANG Y Y, CHEN F D. Screening of salt-tolerance concentration and comparison of salt-tolerance for chrysanthemum and its related taxa. Chinese Journal of Ecology, 2010,29(3):467-472. (in Chinese)
[22]	柏军华, 王克如, 初振东, 陈兵, 李少东. 叶面积测定方法的比较研究. 石河子大学学报(自然科学版), 2005,23(2):89-91.
	BAI J H, WANG K R, CHU Z D, CHEN B, LI S D. Comparative study on the measure methods of the leaf area. Journal of Shihezi University, 2005,23(2):89-91. (in Chinese)
[23]	李志丹, 韩瑞宏, 廖桂兰, 张美华. 植物叶片中叶绿素提取方法的比较研究. 广东第二师范学院学报, 2011,31(3):80-83.
	LI Z D, HAN R H, LIAO G L, ZHANG M H. A comparative study on the different extraction techniques about chlorophyll concentration of plant leaves. Journal of Guangdong University of Education, 2011,31(3):80-83. (in Chinese)
[24]	SU L Y, DAI Z W, LI S H, XIN H P. A novel system for evaluating drough-cold tolerance of grapevines using chlorophyll fluorescence. BMC Plant Biology, 2015,15(1):82. doi: 10.1186/s12870-015-0459-8
[25]	张殿忠, 汪沛洪, 赵会贤. 测定小麦叶片游离脯氨酸的方法. 植物生理学通讯, 1990(4):62-65.
	ZHANG D Z, WANG P H, ZHAO H X. Determination of the content of free proline in wheat leaves. Plant Physiology Communications, 1990(4):62-65. (in Chinese)
[26]	李合生, 孙群, 赵世杰, 章文华. 植物生理实验原理与技术. 北京: 高等教育出版社, 2006: 169-170.
	LI H S, SUN Q, ZHAO S J, ZHANG W H. Principles and Techniques of Plant Physiological Experiment. Beijing: Higher Education Press, 2006: 169-170. (in Chinese)
[27]	王学奎. 植物生理生化实验原理和技术. 北京: 科学出版社, 2006: 167-168, 172-173.
	WANG X K. Principles and Techniques of Plant Physiological Experiment. Beijing: Science Press, 2006: 167-168, 172-173. (in Chinese)
[28]	郑书华, 蔡萍. 电感耦合等离子质谱法测定茶叶中的多种微量元素. 分析仪器, 2018,220(5):40-44.
	ZHENG S H, CAI P. Determination of trace elements in tea by ICP-MS. Analytical Instrumentation, 2018,220(5):40-44. (in Chinese)
[29]	刘德兴, 荆鑫, 焦娟, 魏珉, 隋申利, 赵利华, 李艳玮, 赵娜, 巩彪, 史庆华. 嫁接对番茄产量、品质及耐盐性影响的综合评价. 园艺学报, 2017,44(6):1094-1104.
	LIU D X, JING X, JIAO J, WEI M, SUI S L, ZHAO L H, LI Y W, ZHAO N, GONG B, SHI Q H. Comprehensive evaluation of yield, quality and salt stress tolerance in grafting tomato. Acta Horiculture Sinica, 2017,44(6):1094-1104. (in Chinese)
[30]	李彦, 张英鹏, 孙明, 高弼模. 盐分胁迫对植物的影响及植物耐盐机理研究进展. 中国农学通报, 2008,24(1):258-263.
	LI Y, ZHANG Y P, SUN M, GAO B M. Research advance in the effects of salt stress on plant and the mechanism of plant resistance. Chinese Agricultural Science Bulletin, 2008,24(1):258-263. (in Chinese)
[31]	管志勇, 陈发棣, 滕年军, 陈素梅, 刘浦生. 5种菊花近缘种属植物的耐盐性比较. 中国农业科学, 2010,43(4):787-794.
	GUAN Z Y, CHEN F D, TENG N J, CHEN S M, LIU P S. Study on the NaCl tolerance in five plant species from Dendranthema and its relatives. Scientia Agricultura Sinica, 2010,43(4):787-794. (in Chinese)
[32]	张金林, 李惠茹, 郭姝媛, 王锁民, 施华中, 韩庆庆, 包爱科, 马清. 高等植物适应盐逆境研究进展. 草业学报, 2015,24(12):220-236.
	ZHANG J L, LI H R, GUO S Y, WANG S M, SHI H Z, HAN Q Q, BAO A K, MA Q. Research advances in higher plant adaptation to salt stress. Acta Prataculturae Sinica, 2015,24(12):220-236. (in Chinese)
[33]	COLLA G, ROUPHAEL Y, REA E, CARDARELLI M. Grafting cucumber plants enhance tolerance to sodium chloride and sulfate salinization. Scientia Horticulturae, 2012,135:177-185.
[34]	杨立飞, 朱月林, 胡春梅, 刘正鲁, 张古文. NaCl胁迫对嫁接黄瓜膜脂过氧化、渗透调节物质含量及光合特性的影响. 西北植物学报, 2006,26(6):1195-1200.
	YANG L F, ZHU Y L, HU C M, LIU Z L, ZHANG G W. Effects of NaCl stress on the contents of the substances regulating membrane lipid oxidation and osmosis and photosynthetic characteristics of grafted cucumber. Acta Botanica Boreali-Occidentalia Sinica, 2006,26(6):1195-1200. (in Chinese)
[35]	韩萌. 混合盐碱胁迫对7种禾种子萌发及生理特性的影响[D]. 哈尔滨: 哈尔滨师范大学, 2013.
	HAN M. Effects of saline-alkaloid stress on the seed germination and physiological characteristics of seven grass seeds[D]. Harbin: Harbin Normal University, 2013. (in Chinese)
[36]	陈淑芳, 朱月林, 刘友良, 李式军. NaCl胁迫对番茄嫁接苗保护酶活性、渗透调节物质含量及光合特性的影响. 园艺学报, 2005,32(4):609-613.
	CHEN S F, ZHU Y L, LIU Y L, LI S J. Effects of NaCl stress on activities of protective enzymes, contents of osmotic adjustment substances and photosynthetic characteristics in grafted tomato seedlings. Acta Horiculture Sinica, 2005,32(4):609-613. (in Chinese)
[37]	LLOPEZ-SERRANO L, CANET-SANCHIS G, SELAK G V, PENELLA C, SAN BAUTISTA A, LOPEZ-CALARZA S, CALATAYUD A. Physiological characterization of a pepper hybrid rootstock designed to cope with salinity stress. Plant Physiology and Biochemistry, 2020,148:207-219.
[38]	ZHEN A, BIE Z L, HUANG Y, LIU Z X, LI Q. Effects of scion and rootstock genotypes on the anti-oxidant defense systems of grafted cucumber seedlings under NaCl stress. Soil Science and Plant Nutrition, 2010,56(2):263-271.
[39]	WANG N, QIAO W Q, LIU X H, SHI J B, XU Q H, ZHOU H, YAN G T, HUANG Q. Relative contribution of Na⁺/K⁺, homeostasis, photochemical efficiency and antioxidant defense system to differential salt tolerance in cotton (Gossypium hirsutum L.) cultivars . Plant Physiology and Biochemistry, 2017,119:121-131. pmid: 28866234
[40]	ASHRAF M, FOOLAD M R. Roles of glycine betaine and proline in improving plant abiotic stress resistance. Environmental and Experimental Botany, 2005,59(2):206-216.
[41]	杨凡. 硅肥和虫害及机械损伤对油菜抗虫性的影响[D]. 呼和浩特: 内蒙古农业大学, 2019.
	YANG F. Effects of silicon fertilizer, pests and mechanical damage on insect resistance of rape[D]. Hohot: Inner Mongolia Agricultural University, 2019. (in Chinese)
[42]	MUNNS R, TESTER M. Mechanisms of salinity tolerance. Annual Review of Plant Biology, 2008,59(1):651-681.
[43]	BALIBREA M E, DELL'AMICO J, BOLAR IN M C, PEREZ-ALFOCEA F. Carbon partitioning and sucrose metabolism in tomato plants growing under salinity. Physiologia Plantarum, 2000,110(4):503-511.
[44]	YEO A, FLOWERS T. Plant Solute Transport. Blackwell Publishing Ltd., Oxford, UK, 2007.
[45]	李春燕, 杨廷桢, 王新平, 王淑婷, 王骞, 蔡华成, 杜学梅, 高敬东, 弓桂花. 苹果矮化中间砧对接穗品种枝条输导组织解剖结构的影响. 西北农业学报, 2019,28(3):404-411.
	LI C Y, YANG T Z, WANG X P, WANG S T, WANG Q, CAI H C, DU X M, GAO J D, GONG G H. Effect of apple dwarf rootstock on anatomy structure of conducting tissue in branch of scion. Acta Agriculturae Boreali-occidentalis Sinica, 2019,28(3):404-411. (in Chinese)
[46]	阳燕娟, 郭世荣, 于文进. 嫁接对盐胁迫下西瓜幼苗体内离子和内源激素含量与分布的影响. 西北植物学报, 2015,35(3):80-87.
	YANG Y J, GUO S R, YU W J. Effect of rootstock-grafting on the contents and distributions of ions and endogenous hormones in watermelon seedlings under NaCl stress. Acta Botanica Boreali-Occidentalia Sinica, 2015,35(3):80-87. (in Chinese)
[47]	COBAN A, AKHOUNDNEJAD Y, DERE S, DASGAN H Y. Impact of salt-tolerant rootstock on the enhancement of sensitive tomato plant responses to salinity. Hortscience, 2020,55(1):35-39.
[48]	HUANG Y, BIE Z L, LIU P Y, NIU M L, ZHEN A, LIU Z X, LEI B, GU D G, LU C, WANG B T. Reciprocal grafting between cucumber and pumpkin demonstrates the roles of the rootstock in the determination of cucumber salt tolerance and sodium accumulation. Scientia Horticulturae, 2013,149:47-54.
[49]	王幼群. 植物嫁接系统及其在植物生命科学研究中的应用. 科学通报, 2011,56(30):2478-2485.
	WANG Y Q. Plant grafting and its application in biological research. Chinese Science Bulletin, 2011,56(30):2478-2485. (in Chinese)
[50]	孙敬爽, 李少峰, 董辰希, 孙长忠. 嫁接植物体中RNA分子长距离传递研究进展. 林业科学, 2014,50(11):158-165.
	SUN J S, LI S F, DONG C X, SUN C Z. Research progress and prospect of long distance transmission of RNA molecules in grafted plants. Scientia Silvae Sinicae, 2014,50(11):158-165. (in Chinese)
[51]	孙静宇. 南瓜CmHKT1;1提高黄瓜嫁接苗耐盐性的机理及相关microRNAs鉴定[D]. 武汉: 华中农业大学, 2019.
	SUN J Y. The mechanism of CmHKT1;1 increasing salt tolerance of grafted cucumber seedling and identification of related microRNAs[D]. Wuhan: Huazhong Agricultural University, 2019. (in Chinese)

处理 Treatment
处理 Treatment	1	4	8
自根苗+NaCl Self-rooted seedlings+NaCl	5.30%	7.60%	19.80%
自接苗+NaCl Self-grafted seedlings+NaCl	0	0.35%	6.90%
异根嫁接苗+NaCl Hetero-root grafted seedlings+NaCl	0	0	0.19%

指标 Index	时间 Time (d)	自根苗+CK Self-rooted seedlings+CK	自接苗+CK Self-grafted seedlings+CK	异根嫁接苗+CK Hetero-root grafted seedlings+CK	自根苗+NaCl Self-rooted seedlings+NaCl	自接苗+NaCl Self-grafted seedlings+NaCl	异根嫁接苗+NaCl Hetero-root grafted seedlings+NaCl
超氧化物歧化酶 SOD	0	1497.541c	1723.173b	1999.418a	1565.914c	1749.872b	1926.829a
	1	1515.774ab	1434.051b	1589.519a	1033.416d	1282.002c	1564.286ab
	4	1437.860e	1585.461d	2110.881a	858.959f	1777.534c	1890.630b
	8	1364.903c	1825.189b	2089.685a	1104.757d	1255.950c	1853.107b
过氧化物酶 POD	0	150.400ab	153.667a	147.200b	153.000a	153.100a	149.960ab
	1	148.600c	153.500b	148.920bc	147.467c	159.700a	147.967c
	4	152.833a	151.300ab	148.960ab	141.100c	142.500c	146.567bc
	8	147.633a	148.733a	147.400a	130.700b	127.100b	144.167a
过氧化氢酶 CAT	0	30.360b	35.901a	29.672b	28.773b	31.577b	30.811b
	1	27.826d	40.139c	33.855cd	34.974cd	81.653a	62.958b
	4	50.822c	63.389a	57.141b	33.383d	67.180a	49.207c
	8	58.834ab	65.549a	51.922bc	22.544d	50.527bc	45.853c