固体培养内生真菌对土壤盐胁迫下木麻黄幼苗渗透调节和抗氧化系统的影响

doi:10.17521/cjpe.2022.0061

摘要/Abstract

摘要：

为明晰内生真菌在宿主植物耐盐生长中是否存在拮抗作用, 探讨利用内生真菌提高困难立地条件下造林成活率的可能, 将两株耐盐内生真菌葡萄座腔菌(Botryosphaeria sp.) Z1 (T3)、炭团菌(Hypoxylon sp.) Y6 (T4)及其混合菌(T2)进行固体发酵培养, 添加到栽植有木麻黄(Casuarina equisetifolia)幼苗的不同盐度(质量分数为0、3‰、6‰、9‰)盆钵土壤中, 以添加菌株载体(T1)和无添加(CK)为对照, 研究固体培养内生真菌对土壤盐胁迫下木麻黄幼苗抗逆生理特征的影响及其与生物量的关系。结果表明: 菌株处理显著提高了盐胁迫下幼苗的生物量, 幼苗生理特征受土壤盐度、时间和菌株种类影响较大。3‰盐度下, T2处理15天后可溶性糖(SS)含量、可溶性蛋白(SP)含量、超氧化物歧化酶(SOD)活性以及45天后过氧化氢酶(CAT)活性高于CK、T1, 相对电导率(REC)在T2、T3和T4处理胁迫期都显著降低; 9‰盐度下, T2、T3和T4处理实验期内REC、H₂O₂含量显著降低, 胁迫60天后丙二醛(MDA)含量显著降低, SS含量、SOD活性、CAT活性显著增加, T2处理的SP含量、过氧化物酶(POD)活性同样显著增加。采用逐步回归建立了菌株处理下幼苗生理特征与生物量关系的回归方程模型, 通径分析表明REC、POD活性和SS含量为影响生物量的主要生理因素。综上, 不同菌株对幼苗渗透调节物质含量、氧化酶活性和生物量的影响不同, 混合菌的拮抗作用最为明显, 研究结果初步揭示了盐胁迫下木麻黄渗透调节物质含量、抗逆酶、生物量与木麻黄内生真菌的关系, 为木麻黄抗逆工程菌的发掘提供了研究基础。

关键词: 木麻黄, 土壤盐度, 内生真菌, 植物生理, 生物量, 通径分析

Abstract:

Aims Endophyte is a new microbial resource, which has a complex micro-ecological relationship with plants. In this paper, the antagonistic effect of endophytic fungi on the growth of salt-tolerant plants was confirmed by adding endophytic fungi to the plants and testing the physiological characteristics and biomass of the seedlings.

Methods Two salt-tolerant endophytic fungi Botryosphaeria sp. Z1 (T3), Hypoxylon sp. Y6 (T4), and their mixture (T2) were cultured in solid fermentation and added to potted soil planted with seedlings of Casuarina equisetifolia at different salinity (mass fraction: 0, 3‰, 6‰ and 9‰), which were treated with an added fungi of seedlings. The addition of microbiological medium (T1) and no added microbiological medium (CK) were treated as two experimental controls, which was to confirm the effect of the fungi, but not the microbiological medium. Samples were taken at 15 day intervals, four times, for a total of 60 days, and the relative electronic conductivity (REC), malondialdehyde (MDA) content, soluble sugar (SS) content, soluble protein (SP) content, proline (PRO) content, superoxide dismutase (SOD) activity, peroxidase (POD) activity, catalase (CAT) activity, reactive oxygen (H₂O₂) content of leaves and the biomass of seedling were determined each time.

Important findings We found that the biomass of seedlings treated by fungal strain was significantly increased under salt stress, and the physiological characteristics were significantly affected by soil salinity, time and fungal species. At 3‰ salinity, the contents of SS and SP, and the activity of SOD and CAT were higher in T2 treatment than in CK and T1 after 15 days of treatment, but the REC was significantly lower in T2, T3 and T4 treatments during the stress period. At 9‰ salinity, T2 treatment significantly reduced the REC and the contents of H₂O₂ and MDA, but significantly increased the content of SS and the activity of SOD and CAT. Especially, SP content and POD activity were also significantly increased in T2 treatment after 60 days. A stepwise regression was used to model the relationship between physiological characteristics and biomass of seedlings treated with the strains, and the path analysis showed that REC, POD activity and SS content were the main physiological factors affecting biomass. In conclusion, the effects of different fungi on the content of osmoregulatory substances, oxidase activity and biomass of seedlings were different, and the antagonistic effect of the mixed fungi was the most obviously. Thus, the relationship between osmotic regulatory substances, antioxidant enzymes, biomass and endophytic fungi of C. equisetifolia under salt stress was clarified, which provided a basic for the further study on the stress resistant engineering fungi for C. equisetifolia.

Key words: Casuarina equisetifolia, soil salinity, endophytic fungi, plant physiology, biomass, path analysis

李冠军, 陈珑, 余雯静, 苏亲桂, 吴承祯, 苏军, 李键. 固体培养内生真菌对土壤盐胁迫下木麻黄幼苗渗透调节和抗氧化系统的影响. 植物生态学报, 2023, 47(6): 804-821. DOI: 10.17521/cjpe.2022.0061

LI Guan-Jun, CHEN Long, YU Wen-Jing, SU Qin-Gui, WU Cheng-Zhen, SU Jun, LI Jian. Effects of solid culture endophytic fungi on osmotic adjustment and antioxidant system of Casuarina equisetifolia seedlings under soil salt stress. Chinese Journal of Plant Ecology, 2023, 47(6): 804-821. DOI: 10.17521/cjpe.2022.0061

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2022.0061

https://www.plant-ecology.com/CN/Y2023/V47/I6/804

图/表 11

图1 盐胁迫和菌株处理在不同时间对木麻黄幼苗小枝丙二醛含量和相对电导率的影响(平均值±标准误)。不同小写字母表示在同一盐度同一菌株处理下不同时间差异显著(p < 0.05), 不同大写字母表示同一盐度同一时间不同菌株处理间差异显著(p < 0.05)。CK, 未添加菌株载体; T1, 添加菌株载体; T2, 添加葡萄座腔菌和炭团菌; T3, 添加葡萄座腔菌; T4, 添加炭团菌。

Fig. 1 Effects of salt stress and fungus treatments on malondialdehyde (MDA) content and relatively electronic conductivity (REC) in the branchlets of Casuarina equisitifolia (mean ± SE). Different lowercase letters indicate significant differences between sampling days under the same fungus treatment and the same salinity level (p < 0.05), and different uppercase letters indicate significant differences at different endophytic fungal treatments at the same sampling day (p < 0.05). CK, no added microbiological medium; T1, added microbiological medium; T2, added mixture of Botryosphaeria sp. and Hypoxylon sp.; T3, added Botryosphaeria sp.; T4, added Hypoxylon sp.

图2 盐胁迫和菌株处理在不同时间对木麻黄幼苗小枝可溶性糖含量的影响(平均值±标准误)。不同小写字母表示在同一盐度同一菌株处理下不同时间差异显著(p < 0.05), 不同大写字母表示同一盐度同一时间不同菌株处理间差异显著(p < 0.05)。CK, 未添加菌株载体; T1, 添加菌株载体; T2, 添加葡萄座腔菌和炭团菌; T3, 添加葡萄座腔菌; T4, 添加炭团菌。

Fig. 2 Effects of salt stress and fungus treatments on the content of soluble sugar (SS) in the branchlets of Casuarina equisitifolia (mean ± SE). Different lowercase letters indicate significant differences between sampling days under the same fungus treatment and the same salinity level (p < 0.05), and different uppercase letters indicate significant differences at different endophytic fungal treatments at the same sampling day (p < 0.05). CK, no added microbiological medium; T1, added microbiological medium; T2, added mixture of Botryosphaeria sp. and Hypoxylon sp.; T3, added Botryosphaeria sp.; T4, added Hypoxylon sp.

图3 盐胁迫和菌株处理在不同时间对木麻黄幼苗小枝可溶性蛋白和脯氨酸含量的影响(平均值±标准误)。不同小写字母表示在同一盐度同一菌株处理下不同时间差异显著(p < 0.05), 不同大写字母表示同一盐度同一时间不同菌株处理间差异显著(p < 0.05)。CK, 未添加菌株载体; T1, 添加菌株载体; T2, 添加葡萄座腔菌和炭团菌; T3, 添加葡萄座腔菌; T4, 添加炭团菌。

Fig. 3 Effects of salt stress and fungus treatments on the contents of soluble protein (SP) and proline (PRO) in the branchlets of Casuarina equisitifolia (mean ± SE). Different lowercase letters indicate significant differences between sampling days under the same fungus treatment and the same salinity level (p < 0.05), and different uppercase letters indicate significant differences at different endophytic fungal treatments at the same sampling day (p < 0.05). CK, no added microbiological medium; T1, added microbiological medium; T2, added mixture of Botryosphaeria sp. and Hypoxylon sp.; T3, added Botryosphaeria sp.; T4, added Hypoxylon sp.

图4 盐胁迫和菌株处理在不同时间对木麻黄幼苗小枝超氧化物歧化酶和过氧化物酶活性的影响(平均值±标准误)。不同小写字母表示在同一盐度同一菌株处理下不同时间差异显著(p < 0.05), 不同大写字母表示同一盐度同一时间不同菌株处理间差异显著(p < 0.05)。CK, 未添加菌株载体; T1, 添加菌株载体; T2, 添加葡萄座腔菌和炭团菌; T3, 添加葡萄座腔菌; T4, 添加炭团菌。

Fig. 4 Effects of salt stress and fungus treatments on the activity of superoxide dismutase (SOD) and peroxide (POD) in the branchlets of Casuarina equisitifolia (mean ± SE). Different lowercase letters indicate significant differences between sampling days under the same fungus treatment and the same salinity level (p < 0.05), and different uppercase letters indicate significant differences at different endophytic fungal treatments at the same sampling day (p < 0.05). CK, no added microbiological medium; T1, added microbiological medium; T2, added mixture of Botryosphaeria sp. and Hypoxylon sp.; T3, added Botryosphaeria sp.; T4, added Hypoxylon sp.

图5 盐胁迫和菌株处理在不同时间对木麻黄幼苗小枝过氧化氢酶活性和活性氧含量的影响(平均值±标准误)。不同小写字母表示在同一盐度同一菌株处理下不同时间差异显著(p < 0.05), 不同大写字母表示同一盐度同一时间不同菌株处理间差异显著(p < 0.05)。CK, 未添加菌株载体; T1, 添加菌株载体; T2, 添加葡萄座腔菌和炭团菌; T3, 添加葡萄座腔菌; T4, 添加炭团菌。

Fig. 5 Effects of salt stress and fungus treatments on the activity of catalase (CAT) and hydrogen peroxide (H2O2) in the branchlets of Casuarina equisitifolia at different times (mean ± SE). Different lowercase letters indicate significant differences between sampling days under the same fungus treatment and the same salinity level (p < 0.05), and different uppercase letters indicate significant differences at different endophytic fungal treatments at the same sampling day (p < 0.05). CK, no added microbiological medium; T1, added microbiological medium; T2, added mixture of Botryosphaeria sp. and Hypoxylon sp.; T3, added Botryosphaeria sp.; T4, added Hypoxylon sp.

表1 盐胁迫和菌株处理对木麻黄幼苗生物量的影响(平均值±标准误)

Table 1 Effects of salt stress and fungus treatments on biomass of Casuarina equisitifolia seedlings (mean ± SE)

土壤盐浓度 Soil salinity		地上生物量 Aboveground biomass (g)	地下生物量 Belowground biomass (g)	整株生物量 Total biomass (g)	土壤盐浓度 Soil Salinity		地上生物量 Aboveground biomass (g)	地下生物量 Belowground biomass (g)	整株生物量 Total biomass (g)
0‰	CK	4.37 ± 0.29^a	1.18 ± 0.03^a	5.72 ± 0.33^a	6‰	CK	3.59 ± 0.09^a	0.76 ± 0.01^a	5.58 ± 0.70^a
	T1	4.86 ± 0.54^a	1.36 ± 0.06^a	9.71 ± 0.61^b		T1	3.27 ± 0.07^ab	0.82 ± 0.01^a	6.69 ± 0.71^a
	T2	7.77 ± 0.60^b	3.80 ± 0.08^c	8.97 ± 0.14^b		T2	5.61 ± 0.22^c	1.72 ± 0.03^c	8.33 ± 1.01^a
	T3	8.65 ± 0.16^b	2.05 ± 0.21^b	8.70 ± 0.98^b		T3	4.59 ± 0.66^ab	1.46 ± 0.06^b	5.86 ± 0.85^a
	T4	8.01 ± 1.42^b	2.10 ± 0.22^b	10.11 ± 1.62^b		T4	4.46 ± 0.15^b	1.47 ± 0.03^b	7.22 ± 0.77^a
3‰	CK	4.41 ± 0.19^a	0.97 ± 0.10^a	6.01 ± 0.47^a	9‰	CK	2.18 ± 0.05^a	0.45 ± 0.02^a	4.19 ± 0.19^a
	T1	4.11 ± 0.05^a	1.13 ± 0.01^a	6.03 ± 0.35^a		T1	2.62 ± 0.04^b	0.61 ± 0.01^b	5.33 ± 0.33^a
	T2	6.55 ± 0.12^b	2.55 ± 0.22^c	8.10 ± 0.49^b		T2	4.77 ± 0.09^d	1.52 ± 0.01^d	7.35 ± 0.87^b
	T3	6.05 ± 0.68^b	1.78 ± 0.18^b	7.67 ± 0.87^ab		T3	4.78 ± 0.14^d	1.30 ± 0.08^c	7.46 ± 0.79^b
	T4	5.50 ± 0.11^b	1.76 ± 0.07^b	6.09 ± 0.32^b		T4	3.06 ± 0.04^c	1.29 ± 0.04^c	8.00 ± 0.57^b

表2 土壤盐胁迫下木麻黄幼苗生理特征与生物量的相关系数

Table 2 Correlation coefticients between biomass and physiologieal characteristics of Casuarina equisitifolia seedlings under soil salt stress

表3 菌株处理木麻黄幼苗生理特征与生物量间的相关系数

Table 3 Correlation coefficients between biomass and physiological characteristics of Casuarina equisitifolia seedlings under fungal strain treatment

	丙二醛含量 Malondialdehyde content	超氧化物歧化酶活性 Superoxide dismutase activity	过氧化物酶活性 Peroxide activity	活性氧含量 Hydrogen peroxide content	过氧化氢酶活性 Catalase activity	可溶性糖含量 Soluble sugar content	可溶性蛋白含量 Soluble protein content	脯氨酸含量 Proline content	相对电导率 Relatively conductivity
地上生物量 Aboveground biomass	-0.448^**	0.311	-0.703^**	-0.166	-0.664^**	-0.718^**	-0.154	-0.094	-0.584^**
地下生物量 Belowground biomass	-0.330^*	0.252	-0.650^**	-0.263	-0.527^**	-0.642^**	-0.239	-0.023	-0.497^**
整株生物量 Total biomass	-0.410^*	0.466^**	-0.805^**	-0.237	-0.659^**	-0.725^**	-0.159	-0.079	-0.497^**

表4 土壤盐胁迫下木麻黄幼苗生理特征与生物量的多元逐步回归分析

Table 4 Multiple stepwise regression analysis of biomass and physiological characteristics of Casuarina equisitifolia seedlings under soil salinity

土壤盐度 Soil salinity	菌株处理 Endophytic fungi	生物量 Biomass	主要影响因子 Major affecting factor	回归方程 Regression equation	R²
3‰	T2	地上生物量 Aboveground biomass	SP	Y₁ = 13.727SP	0.998^**
		地下生物量 Belowground biomass	SP	Y₂ = 4.165SP	0.999^**
		整株生物量 Total biomass	SP	Y₃ = 17.892SP	0.998^**
	T3	地上生物量 Aboveground biomass	CAT	Y₁ = 0.071CAT	0.997^**
		地下生物量 Belowground biomass	H₂O₂	Y₂ = 1.584H₂O₂	0.995^**
		整株生物量 Total biomass	CAT	Y₃ = 0.902CAT	0.997^**
	T4	地上生物量 Aboveground biomass	REC, PRO, SOD	Y₁ = 21.573REC - 0.528PRO - 0.001SOD	0.998^**
		地下生物量 Belowground biomass	PRO	Y₂ = 1.069PRO	0.978^**
		整株生物量 Total biomass	REC	Y₃ = 24.055REC	0.999^**
6‰	T2	地上生物量 Aboveground biomass	SP	Y₁ = 15.411SP	0.992^**
		地下生物量 Belowground biomass	REC	Y₂ = 4.748REC	0.994^**
		整株生物量 Total biomass	SS	Y₃ = 0.668SS	0.993^**
	T3	地上生物量 Aboveground biomass	MDA	Y₁ = 0.245MDA	0.992^**
		地下生物量 Belowground biomass	MDA, CAT, SS	Y₂ = 0.099MDA - 0.007CAT + 0.005SS	0.991^**
		整株生物量 Total biomass	MDA	Y₃ = 0.313MDA	0.993^**
	T4	地上生物量 Aboveground biomass	H₂O₂	Y₁ = 5.133H₂O₂	0.996^**
		地下生物量 Belowground biomass	SS, PRO, CAT	Y₂ = 0.419SS - 2.023PRO + 0.001CAT	0.995^**
		整株生物量 Total biomass	H₂O₂	Y₃ = 6.777H₂O₂	0.995^**
9‰	T2	地上生物量 Aboveground biomass	SOD, SP, POD	Y₁ = 0.061SOD - 19.377SP - 0.004POD	0.967^**
		地下生物量 Belowground biomass	CAT, REC, SP	Y₂ = 0.059CAT - 18.461REC - 0.102SP	0.973^**
		整株生物量 Total biomass	SOD	Y₃ = 0.026SOD	0.991^**
	T3	地上生物量 Aboveground biomass	REC, POD, PRO	Y₁ = 36.719REC - 0.038POD - 0.945PRO	0.970^**
		地下生物量 Belowground biomass	MDA, SS, SOD	Y₂ = 0.212MDA - 0.085SS - 0.001SOD	0.962^**
		整株生物量 Total biomass	SP, MDA, PRO	Y₃ = 35.105SP - 0.391MDA + 0.065PRO	0.991^**
	T4	地上生物量 Aboveground biomass	SOD	Y₁ = 0.028SOD	0.985^**
		地下生物量 Belowground biomass	MDA	Y₂ = 0.135MDA	0.972^**
		整株生物量 Total biomass	POD, H₂O₂, SS	Y₃ = 0.419POD - 45.734H₂O₂ - 0.015SS	0.985^**

表4 土壤盐胁迫下木麻黄幼苗生理特征与生物量的多元逐步回归分析

Table 4 Multiple stepwise regression analysis of biomass and physiological characteristics of Casuarina equisitifolia seedlings under soil salinity

土壤盐度 Soil salinity	菌株处理 Endophytic fungi	生物量 Biomass	主要影响因子 Major affecting factor	回归方程 Regression equation	R²
3‰	T2	地上生物量 Aboveground biomass	SP	Y₁ = 13.727SP	0.998^**
		地下生物量 Belowground biomass	SP	Y₂ = 4.165SP	0.999^**
		整株生物量 Total biomass	SP	Y₃ = 17.892SP	0.998^**
	T3	地上生物量 Aboveground biomass	CAT	Y₁ = 0.071CAT	0.997^**
		地下生物量 Belowground biomass	H₂O₂	Y₂ = 1.584H₂O₂	0.995^**
		整株生物量 Total biomass	CAT	Y₃ = 0.902CAT	0.997^**
	T4	地上生物量 Aboveground biomass	REC, PRO, SOD	Y₁ = 21.573REC - 0.528PRO - 0.001SOD	0.998^**
		地下生物量 Belowground biomass	PRO	Y₂ = 1.069PRO	0.978^**
		整株生物量 Total biomass	REC	Y₃ = 24.055REC	0.999^**
6‰	T2	地上生物量 Aboveground biomass	SP	Y₁ = 15.411SP	0.992^**
		地下生物量 Belowground biomass	REC	Y₂ = 4.748REC	0.994^**
		整株生物量 Total biomass	SS	Y₃ = 0.668SS	0.993^**
	T3	地上生物量 Aboveground biomass	MDA	Y₁ = 0.245MDA	0.992^**
		地下生物量 Belowground biomass	MDA, CAT, SS	Y₂ = 0.099MDA - 0.007CAT + 0.005SS	0.991^**
		整株生物量 Total biomass	MDA	Y₃ = 0.313MDA	0.993^**
	T4	地上生物量 Aboveground biomass	H₂O₂	Y₁ = 5.133H₂O₂	0.996^**
		地下生物量 Belowground biomass	SS, PRO, CAT	Y₂ = 0.419SS - 2.023PRO + 0.001CAT	0.995^**
		整株生物量 Total biomass	H₂O₂	Y₃ = 6.777H₂O₂	0.995^**
9‰	T2	地上生物量 Aboveground biomass	SOD, SP, POD	Y₁ = 0.061SOD - 19.377SP - 0.004POD	0.967^**
		地下生物量 Belowground biomass	CAT, REC, SP	Y₂ = 0.059CAT - 18.461REC - 0.102SP	0.973^**
		整株生物量 Total biomass	SOD	Y₃ = 0.026SOD	0.991^**
	T3	地上生物量 Aboveground biomass	REC, POD, PRO	Y₁ = 36.719REC - 0.038POD - 0.945PRO	0.970^**
		地下生物量 Belowground biomass	MDA, SS, SOD	Y₂ = 0.212MDA - 0.085SS - 0.001SOD	0.962^**
		整株生物量 Total biomass	SP, MDA, PRO	Y₃ = 35.105SP - 0.391MDA + 0.065PRO	0.991^**
	T4	地上生物量 Aboveground biomass	SOD	Y₁ = 0.028SOD	0.985^**
		地下生物量 Belowground biomass	MDA	Y₂ = 0.135MDA	0.972^**
		整株生物量 Total biomass	POD, H₂O₂, SS	Y₃ = 0.419POD - 45.734H₂O₂ - 0.015SS	0.985^**

表5 菌株处理木麻黄幼苗生理特征与生物量的多元逐步回归分析

Table 5 Multiple step-up regression analysis of biomass and physiological characteristics of Casuarina equisitifolia seedlings under fungal strain treatment

项目 Item	主要影响因子 Major affecting factor	回归方程 Regression equation	R²
地上生物量 Aboveground biomass	REC, SS, POD	Y₁ = 0.140SS - 14.390REC - 0.017POD	0.621^*
地下生物量 Belowground biomass	POD, REC	Y₂ = 0.009POD - 5.460REC	0.482^*
整株生物量 Total biomass	POD, REC	Y₃ = 0.046POD - 17.826REC	0.691^*

表6 菌株处理下木麻黄幼苗生理特征对生物量影响的通径系数

Table 6 Path coefficients between physiological characteristics and biomass of Casuarina equisitifolia seedlings under fungal strain treatment

因变量 Dependent variable	自变量 Independent variable	直接通径系数 Direct path coefficient	间接通径系数 Indirect path coefficient				剩余通径系数 Residual path coefficient
因变量 Dependent variable	自变量 Independent variable	直接通径系数 Direct path coefficient	SS	REC	POD	合计 Total	剩余通径系数 Residual path coefficient
地上生物量 Aboveground biomass	SS	-0.239	-	-0.165	-0.315	-0.479	0.616
	REC	-0.329	-0.119	-	-0.136	-0.255
	POD	-0.412	-	-0.182	-0.108	-0.291
地下生物量 Belowground biomass	REC	-0.317	-	-	-0.180	-0.180	0.720
地下生物量 Belowground biomass	POD	-0.545	-	-0.104	-	-0.104
整株生物量 Total biomass	REC	-0.260	-	-	-0.237	-0.237	0.556
整株生物量 Total biomass	POD	-0.720	-	-0.085	-	-0.085

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