盐胁迫条件下哈茨木霉ST02对椒样薄荷生长及根区土壤理化性质的影响

doi:10.13560/j.cnki.biotech.bull.1985.2021-1251

摘要/Abstract

摘要：

为研究耐盐木霉对植物耐盐性及盐渍土壤改良的作用,以耐盐哈茨木霉ST02和耐盐椒样薄荷为研究对象,分析盐胁迫条件下ST02对椒样薄荷生理生长、椒样薄荷根区土壤理化性质和土壤中酶活性大小的影响;并通过椒样薄荷生理生长与土壤因子相关性分析和主成分分析,分析影响椒样薄荷生理生长的主要土壤因子。结果表明,ST02缓解盐胁迫对椒样薄荷生长和光合作用的抑制,促进盐胁迫条件下椒样薄荷的生长和光合作用强度;NaCl胁迫条件下ST02减轻椒样薄荷细胞中丙二醛（malondialdehyde,MDA）、可溶性蛋白（soluble protein,SP）和脯氨酸（proline,Pro）的积累,增强超氧化物歧化酶（superoxide dismutase,SOD）、过氧化物酶（peroxidase,POD）和过氧化氢酶（catalase,CAT）活性,通过抗氧化防御途径增强椒样薄荷耐盐性;NaCl胁迫条件下,椒样薄荷生理生长综合指数（physiological/growth comprehensive index,PGCI）为-2.391,ST02处理促进椒样薄荷生理生长（指数为-0.025）;NaCl胁迫条件下,ST02能影响土壤理化性质,降低根区土壤盐浓度（降低17.36%）,并显著提高土壤中碱解氮（alkali-hydrolyzed nitrogen,AN）的含量;ST02能够影响NaCl胁迫条件下椒样薄荷根区土壤酶活性,显著提高土壤中纤维素酶（soil cellulase,SCL）和过氧化氢酶（soil catalase,SCAT）活性;通过椒样薄荷根区土壤理化性质与椒样薄荷生理生长的相关性分析,表明土壤理化性质和土壤酶活性影响椒样薄荷的生长与生理性状;主成分分析影响椒样薄荷生理生长最主要的土壤因子为：EC（electrical conductance）>SCL>SSC（soil saccharase）/SNEP（soil neutrality phosphatase）>AP（available phosphorus）>pH。

关键词: 耐盐木霉, 耐盐植物, 椒样薄荷, 生理生长指数, 土壤理化性质, 土壤酶活性

Abstract:

The salt-resistant Trichoderma harzianum strain ST02 and salt-resistant peppermint were used in this study to analyze the effects of Trichoderma on plant salt resistance and saline soil improvement. The physiological and growth traits of peppermint,the soil physicochemical property and soil enzyme activity of root zone soil under salt stress were analyzed. And the main soil factors related to the physiological and growth of peppermint were analyzed through the correlation analysis and principal component analysis（PCA）. The results showed ST02 relieved the inhibition of NaCl on peppermint growth and photosynthesis and promoted its growth and photosynthesis intensity under salt stress. When treated with NaCl,ST02 reduced the accumulation of maldehyde aldehyde（MDA）,soluble protein（SP）and proline（Pro）in peppermint cells,enhanced superoxide dismutase（SOD）,peroxide dismutase（POD）,and catalase（CAT）activities,and enhanced the salt resistance of peppermint through antioxidant response. The physiological/growth comprehensive index（PGCI）of peppermint treated with NaCl was -2.391,ST02 promoted the physiological and growth of peppermint,as index was -0.025. ST02 influenced the soil physicochemical properties for root zone soil of peppermint under NaCl stress,reduced salinity by 17.36%,and significantly enhanced the content of alkali-hydrolyzed nitrogen（AN）. The soil enzyme activity for the root zone soil of peppermint was also affected by ST02 strain,and the activity of soil cellulase（SCL）and catalase（SCAT）increased remarkably. The soil physicochemical property and soil enzyme activity affected the physiological and growth traits under NaCl stress through the correlation analysis. And the main soil factor for physiological and growth of peppermint was：EC（electrical conductance）> SCL >SSC（soil saccharase）/SNEP（soil neutrality phosphatase）>AP（available phosphorus）>pH.

Key words: salt-resistant Trichoderma, salt-resistant plant, Mentha×piperita L., physiological and growth index, soil physicochemical property, soil enzyme activity

赵忠娟, 杨凯, 扈进冬, 魏艳丽, 李玲, 徐维生, 李纪顺. 盐胁迫条件下哈茨木霉ST02对椒样薄荷生长及根区土壤理化性质的影响[J]. 生物技术通报, 2022, 38(7): 224-235.

ZHAO Zhong-juan, YANG Kai, HU Jin-dong, WEI Yan-li, LI Ling, XU Wei-sheng, LI Ji-shun. Effects of Trichoderma harzianum ST02 on the Growth of Peppermint and Physicochemical Properties of Root Zone Soil Under Salt Stress[J]. Biotechnology Bulletin, 2022, 38(7): 224-235.

图/表 12

表1 实验设计分组及处理方法

Table 1 Experimental grouping and processing method

分组Group		处理方法Processing method
实验组 Experimental group	NaCl处理 NaCl treatment	椒样薄荷茎段扦插7 d后,用200 mL水浇灌;再7 d后,用200 mmol/L NaCl溶液200 mL浇灌,每隔3 d 进行1次NaCl处理,连续浇灌3次后用正常水浇灌
	ST02处理 ST02 treatment	椒样薄荷茎段扦插7 d后,将ST02菌剂用水按1∶100（V∶V）比例混匀（活菌数2×10⁷ CFU/g）制成菌液,用200 mL菌液进行浇灌;ST02处理7 d后,NaCl处理组用NaCl浇灌的同一时间,用正常水浇灌
	NaCl+ST02处理 NaCl+ST02 treatment	椒样薄荷茎段扦插7 d后,将ST02菌剂用水按1∶100（V∶V）比例混匀（活菌数2×10⁷ CFU/g）制成菌液,用200 mL菌液进行浇灌;ST02处理7 d后,用200 mmol/L NaCl溶液200 mL浇灌,每隔3 d进行1次NaCl处理,连续浇灌3次后用正常水浇灌
对照组Control		实验组用ST02菌液和NaCl处理的同时,用200 mL水浇灌

图1 不同处理对椒样薄荷生长和光合作用的影响 A：椒样薄荷的株高;B：椒样薄荷地上部分的鲜重;C：椒样薄荷净光合速率（Pn）;D：椒样薄荷叶片中叶绿素含量（CK为对照组,+NaCl为NaCl处理实验组,+ST02为哈茨木霉ST02处理实验组,+NaCl+ST02为NaCl和ST02联合处理实验组,数值为平均值±标准差（n=5）,不同小写字母表示组间ANOVA方差分析,P<0.05表示存在显著性差异,下同

Fig.1 Growth and photosynthesis intensity of peppermint under different treatments A：The stem length of peppermint（Mentha×piperita L）. B：The fresh weight of peppermint aboveground parts. C：The Pn value of peppermint. D：The chlorophyll content of peppermint leaves.（CK：control check;+NaCl：NaCl treatment;+ST02：ST02 treatment;+NaCl+ST02：NaCl and ST02 combined treatment. The bars are standard error of the mean（n=5）,different letters were significantly different according to ANOVA analysis（P<0.05）, the same below

图2 不同处理对椒样薄荷氧化损伤和渗透调节相关物质含量的影响 A：椒样薄荷叶片丙二醛（MDA）含量;B：椒样薄荷叶片可溶性蛋白（SP）含量;C：椒样薄荷叶片脯氨酸（Pro）含量

Fig.2 Oxidative damage and osmotic regulatory substances contents of peppermint under different treatments A：The malondialdehyde（MDA）content of peppermint leaves. B：The content of soluble protein（SP）in peppermint leaves. C：The content of proline（Pro）in peppermint leaves

图3 不同处理对椒样薄荷抗氧化酶活性的影响 A ：椒样薄荷叶片丙二醛（MDA）含量；B ：椒样薄荷叶片可溶性蛋白（SP）含量；C ：椒样薄荷叶片脯氨酸（Pro）含量

Fig.3 Influences of different treatments on peppermint antioxidase activity A：The activity of superoxide dismutase（SOD）in peppermint leaves. B：The activity of peroxidase（POD）in peppermint leaves. C：The activity of catalase（CAT）in peppermint leaves

表2 不同处理椒样薄荷生理生长综合指数

Table 2 Physiological/growth comprehensive index（PGCI）of peppermint under different treatments

不同处理 Treatment	PGI1 SL	PGI2 FW	PGI3 Pn	PGI4 SPAD	PGI5 SOD	PGI6 POD	PGI7 CAT	PGCI
NaCl	-0.154	-0.155	-0.363	-0.371	-0.481	-0.391	-0.476	-2.391
ST02	0.099	0.056	0.014	0.040	0.146	0.212	0.176	0.743
NaCl+ST02	0.007	-0.024	0.011	0.037	-0.146	0.061	0.029	-0.025

表3 不同处理椒样薄荷根区土壤理化性质

Table 3 Physicochemical properties of peppermint root zone soil under different treatments

不同处理 Treatment	pH	EC/（μs·cm^-1）	CEC/（cmol·kg^-1）	OM/（g·kg^-1）	AN/（mg·kg^-1）	AP/（mg·kg^-1）	AK/（mg·kg^-1）
CK	7.56±0.09^b	182.80±6.02^c	16.13±2.01^b	20.04±3.5^a	99.70±21.69^a,b	26.84±9.02^a	140.22±17.64^a
NaCl	7.66±0.06^a	406.54±35.10^a	16.72±3.32^b	17.00±4.7^a	87.62±13.83^b	19.38±4.58^a	125.20±15.53^a
ST02	7.56±0.05^b	183.28±3.14^c	21.98±3.53^a	21.48±5.9^a	119.05±23.33^a,b	28.78±5.22^a	126.88±14.32^a
NaCl+ST02	7.68±0.06^a	335.98±49.07^b	18.28±0.98^b	22.50±6.0^a	128.60±30.96^a	27.18±9.73^a	125.85±14.74^a

图4 不同处理对椒样薄荷根区土壤酶活性的影响 A：土壤脲酶（SUE）活性;B：土壤蔗糖酶（SSC）活性;C：土壤纤维素酶（SCL）活性;D：土壤磷酸酶（SNEP）活性;E：土壤过氧化氢酶（SCAT）活性

Fig. 4 Influences of different treatments on the activities of soil enzyme A：The activity of soil urease. B：The activity of soil saccharase. C：The activity of soil cellulose. D：The activity of soil neutrality phosphatase. E：The activity of soil catalase

图5 椒样薄荷生理生长和根际土壤理化性质相关性英文缩写注释同上

Fig.5 Correlation analysis between physiological/growth index of peppermint and physicochemical properties of its root zone soil Abbreviated notes are the same as above

表4 椒样薄荷生理生长性状和根际土壤理化性质相关系数

Table 4 Correlation coefficient between physiological/growth indexes of peppermint and physicochemical properties of its root zone soil

指标Index	pH	EC	CEC	OM	AN	AP	AK	SUE	SSC	SCL	SNEP	SCAT
SL	-0.528^*	-0.719^**	0.398	0.282	0.413	0.28	-0.012	-0.427	0.359	0.644*	0.424	0.084
FW	-0.253	-0.735**	0.546*	0.669**	0.333	0.562**	0.334	0.358	0.478*	0.574**	0.451*	0.065
Pn	-0.355	-0.619**	0.244	0.299	0.196	0.340	0.279	0.020	0.331	0.397	0.441	0.167
SPAD	-0.140	-0.596**	0.284	0.377	0.410	0.271	0.021	0.237	0.262	0.663**	0.192	-0.029
MDA	0.468*	0.785**	-0.332	-0.312	-0.373	-0.377	-0.214	-0.374	-0.523*	-0.568**	-0.355	-0.021
SP	0.500*	0.844**	-0.209	-0.241	-0.326	-0.433	-0.116	-0.309	-0.537*	-0.514*	-0.317	0.029
Pro	0.560*	0.882**	-0.356	-0.256	-0.340	-0.454*	-0.155	-0.319	-0.617**	-0.454*	-0.427	0.113
SOD	-0.487*	-0.884**	0.408	0.300	0.373	0.464*	0.110	0.473*	0.558*	0.549*	0.578**	-0.074
POD	-0.421	-0.758**	0.548*	0.408	0.424	0.535*	0.131	0.358	0.333	0.706**	0.510*	0.148
CAT	-0.445*	-0.775**	0.453*	0.420	0.473	0.481*	0.248	0.422	0.440	0.619**	0.389	0.121
SUE	-0.214	-0.323	0.480*	0.000	0.445*	0.050	-0.167	1	0.220	0.215	0.508*	-0.058
SSC	-0.669**	-0.813**	-0.020	0.151	-0.191	0.229	0.388	0.220	1	-0.178	0.302	-0.553*
SCL	-0.094	-0.215	0.327	0.413	0.560*	0.433	-0.103	0.215	-0.178	1	0.195	0.534*
SNEP	-0.374	-0.505*	0.648**	0.100	0.091	0.222	0.121	0.508*	0.302	0.195	1	-0.083
SCAT	0.161	0.330	0.161	0.052	0.476*	0.257	0.137	-0.058	-0.553*	0.534*	-0.083	1

表5 各指标主成分分析

Table 5 Principal component analysis

成分 Component	特征根 Characteristic root	方差贡献率 Variance contribution ratio/%	累计贡献率 Accumulated variance contribution rate/%
1	10.949	49.766	49.766
2	2.991	13.595	63.361
3	1.929	8.770	72.131
4	1.635	7.430	79.561
5	1.092	4.962	84.523

图6 椒样薄荷生长生理和土壤理化性质PCA分析 A：不同处理样品主成分分析得分图;B：不同指标对主成分分析因子载荷系数图

Fig.6 Principal component analysis of peppermint physiological/growth index and physicochemical properties of root zone soil A：The score for principal component analysis. B：The loading factor of principal component analysis

表6 各指标主成分分析后成分载荷矩阵与贡献率

Table 6 Loading factors and contribution ratio of principal component analysis for each index

椒样薄荷生长生理指标 Physiological /growth index of peppermint	主成分载荷Loading factors		土壤理化性质 Physicochemical properties of root zone soil	主成分载荷Loading factor
椒样薄荷生长生理指标 Physiological /growth index of peppermint	PC1	PC2	土壤理化性质 Physicochemical properties of root zone soil	PC1	PC2
SL	0.892	0.084	pH	-0.508	0.507
FW	0.879	0.079	EC	-0.751	0.579
Pn	0.752	0.004	SCEC	0.491	0.291
SPAD	0.789	0.196	OM	0.418	0.289
MDA	-0.938	0.021	AN	0.407	0.610
SP	-0.913	0.090	AP	0.513	0.159
Pro	-0.931	0.166	AK	0.223	-0.228
SOD	0.963	-0.087	SUE	0.439	0.022
POD	0.959	0.165	SSC	0.535	-0.777
CAT	0.953	0.088	SCL	0.611	0.664
			SNEP	0.535	-0.153
			SCAT	0.037	0.763

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