内生真菌感染对宿主羽茅及邻生植物抗病性的影响

doi:10.17521/cjpe.2021.0132

摘要/Abstract

摘要：

Epichloë内生真菌能够影响宿主植物的抗病性, 但目前关于内生真菌感染对宿主邻生植物抗病性影响的研究很少。该研究首先调查了内蒙古呼伦贝尔天然草地中植物病害发生情况, 发现在常见牧草中禾本科植物病害最为严重, 其中感染内生真菌的羽茅(Achnatherum sibiricum)的总病情指数最低。进一步比较了与羽茅邻生及非邻生植物的病害差异, 结果显示羽茅显著降低了邻生羊草(Leymus chinensis)、狼针草(Stipa baicalensis)和冰草(Agropyron cristatum)的褐斑病病情指数。为探究羽茅对邻生植物病害的影响是否与其内生真菌有关, 分别从内生真菌发酵液、离体叶片和植株层面进行了室内验证试验。结果表明: 羽茅的2种内生真菌Epichloë gansuensis和Epichloë sibiricum发酵液显著抑制了新月弯孢(Curvularia lunata)、根腐离蠕孢(Bipolaris sorokiniana)、核盘菌(Sclerotinia sclerotioru)和三叶草核盘菌(Sclerotinia trifoliorum)的生长; 对于离体叶片, E. gansuensis和E. sibiricum显著减小了宿主叶片受4种病原真菌侵染后的病斑长度。在新月弯孢和根腐离蠕孢的植株接种条件下, 内生真菌显著降低了宿主羽茅及邻生植物羊草的病害程度。该研究首次发现并验证了羽茅内生真菌对邻生禾草抵御褐斑病有一定的增益作用。

关键词: 内生真菌, 羽茅, 邻生禾草, 病原真菌, 褐斑病

Abstract:

Aims Epichloë infection can affect the fungal disease resistance of host grasses. However, few studies have been reported on the effects of endophyte infection on non-symbiotic neighbours.
Methods We surveyed the plant diseases in natural grassland, and compared differences of total disease index between neighbor and non-neighbor plants of Achnatherum sibiricum. Then the laboratory experiments were conducted to investigate the effects of endophyte on the growth of four pathogen species as well as the brown patch of the host and its neighbor plants.
Important findings Major epidemic diseases of grasses were spot blight, rust disease and powdery mildew in Hulun Buir natural grassland. Among common herbages, the total disease index of endophyte-infected A. sibiricum was the lowest. Compared with non-neighbor plants, the brown patch disease index of Leymus chinensis, Stipa baicalensis and Agropyron cristatum was significantly reduced when neighbouring with A. sibiricum. The laboratory experiments results showed that the culture filtration of both Epichloë gansuensis and Epichloë sibiricum significantly restrained the growth of Curvularia lunata, Bipolaris sorokiniana, Sclerotinia sclerotioru and Sclerotinia trifoliorum. The two species of endophytes could reduce lesion area of detached host leaves. In the intact leaves, the endophyte reduced the disease resistance of both the host and its neighbor grasses L. chinensis to C. lunata and B. sorokiniana. This study first verified that the endophytes in A. sibiricum have a positive effect on disease resistance of neighbor grasses to brown patch.

Key words: endophyte, Achnatherum sibiricum, neighbor grasses, fungal pathogens, brown patch

石新建, 张靖歆, 秦天姿, 刘金铭, 高玉葆, 任安芝. 内生真菌感染对宿主羽茅及邻生植物抗病性的影响. 植物生态学报, 2021, 45(8): 860-869. DOI: 10.17521/cjpe.2021.0132

SHI Xin-Jian, ZHANG Jing-Xin, QIN Tian-Zi, LIU Jin-Ming, GAO Yu-Bao, REN An-Zhi. Effects of endophyte infection on fungal disease resistance of Achnatherum sibiricum and non-symbiotic neighbours. Chinese Journal of Plant Ecology, 2021, 45(8): 860-869. DOI: 10.17521/cjpe.2021.0132

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2021.0132

https://www.plant-ecology.com/CN/Y2021/V45/I8/860

图/表 8

表1 呼伦贝尔草原植物病害

Table 1 Plant diseases in Hulun Buir grassland

科 Family	属数 Genus number	物种数 Species number	草原病情指数 Grassland diseases index
科 Family	属数 Genus number	物种数 Species number	斑枯病 Spot blight	黄斑病 Yellow blotch	锈病 Rust disease	褐斑病 Brown patch	白粉病 Powdery mildew	其他病害 Others	累计 Total
禾本科 Gramineae	10	10	13.0	6.4	8.0	5.5	6.8	0.2	39.8
豆科 Leguminosae	5	6	22.8	0.5	2.2	3.9	4.3	-	33.7
菊科 Compositae	7	14	17.4	3.0	1.2	1.9	8.6	0.6	32.7
蔷薇科 Rosaceae	2	6	21.0	7.7	5.0	1.0	1.0	-	35.8
毛茛科 Ranunculaceae	3	5	21.2	5.0	5.1	1.8	0.9	1.0	35.0
百合科 Liliaceae	2	3	34.7	8.1	3.3	1.2	-	0.2	47.4
鸢尾科 Iridaceae	1	3	29.7	17.8	1.4	1.0	0.2	-	50.1
唇形科 Labiatae	2	2	34.4	-	-	-	-	-	34.4
伞形科 Umbelliferae	2	2	20.9	1.2	-	0.1	-	0.2	22.3
石竹科 Caryophyllaceae	2	2	5.4	14.2	-	-	25.0	-	44.6
车前科 Plantaginaceae	1	1	20.0	-	-	-	46.7	-	66.7
川续断科 Dipsacaceae	1	1	40.0	11.7	-	-	8.9	-	60.6
大戟科 Euphorbiaceae	1	1	15.0	-	-	-	-	-	15.0
景天科 Crassulaceae	1	1	-	-	-	-	-	-	0
桔梗科 Campanulaceae	1	1	18.5	2.8	-	0.3	-	0.6	22.2
蓼科 Polygonaceae	1	1	22.5	41.3	-	-	-	-	63.8
柳叶菜科 Onagraceae	1	1	30.0	22.5	-	10.0	-	-	62.5
龙胆科 Gentianaceae	1	1	7.5	15.0	-	-	16.7	-	39.2
马钱科 Loganiaceae	1	1	48.3	-	22.9	-	-	-	71.2
茜草科 Rubiaceae	1	1	11.4	4.1	0.2	1.0	-	-	16.7
莎草科 Cyperaceae	1	1	49.7	1.1	2.6	1.4	0.1	-	54.9
十字花科 Brassicaceae	1	1	7.5	11.7	-	-	7.8	-	27.0
苋科 Amaranthaceae	1	1	-	-	-	-	-	-	0
玄参科 Scrophulariaceae	1	1	-	-	-	3.8	13.3	-	17.1
旋花科 Convolvulaceae	1	1	-	22.5	-	-	-	-	22.5

图1 呼伦贝尔草原常见牧草的总病情指数。Ac, 冰草; As, 羽茅; Cp, 柄状薹草; Il, 马蔺; Lc, 羊草; Pc, 白头翁; Sb, 狼针草; Sd, 防风; Ts, 展枝唐松草; Vs, 野豌豆。

Fig. 1 Total disease index of common herbages in Hulun Buir grassland. Ac, Agropyron cristatum; As, Achnatherum sibiricum; Cp, Carex pediformis; Il, Iris lactea; Lc, Leymus chinensis; Pc, Pulsatilla chinensis; Sb, Stipa baicalensis; Sd, Saposhnikovia divaricata; Ts, Thalictrum squarrosum; Vs, Vicia sepium.

图2 呼伦贝尔草原天然草地中与羽茅邻生和非邻生常见禾草的病情指数分布。A, D, 羊草。B, E, 狼针草。C, F, 冰草。p经非参数Mann-Whitney U检验, p < 0.05表明邻生与非邻生差异显著。

Fig. 2 Frequency distribution of disease index of neighbor and non-neighbor plants with Achnatherum sibiricum of the native grassland in Hulun Buir grassland. A, D, Leymus chinensis. B, E, Stipa baicalensis. C, F, Agropyron cristatum. p was tested by nonparametric Mann-Whitney U test, p < 0.05 indicate significant difference between neighbor and non-neighber.

图3 不同内生真菌发酵液对4种病原真菌的平均抑制率(平均值±标准误, n = 5)。不同大写字母表示病原真菌间差异显著(p < 0.05)。EG, Epichloë gansuensis; ES, Epichloë sibiricum。Bs, 根腐离蠕孢; Cl, 新月弯孢; Ss, 核盘菌; St, 三叶草核盘菌。

Fig. 3 Mean inhibition rate of culture filtrate of different endophytes on four species of pathogenic fungi (mean ± SE, n = 5). Different uppercase letters indicate significant difference among pathogenic fungis (p < 0.05). EG, Epichloë gansuensis; ES, Epichloë sibiricum. Bs, Bipolaris sorokiniana; Cl, Curvularia lunata; Ss, Sclerotinia sclerotioru; St, Sclerotinia trifoliorum.

图4 感染不同内生真菌的羽茅离体叶片对病原真菌侵染后的平均病斑抑制率(平均值±标准误, n = 5)。不同大写字母表示不同病原真菌间差异显著(p < 0.05)。EF为无内生真菌感染的羽茅叶片; EG为感染Epichloë gansuensis的羽茅叶片; ES为感染Epichloë sibiricum的羽茅叶片。Bs, 根腐离蠕孢; Cl, 新月弯孢; Ss, 核盘菌; St, 三叶草核盘菌。

Fig. 4 Mean inhibition rate of disease area on detached leaves of Achnatherum sibiricum after inoculation with 4 pathogenic fungi (means ± SE, n = 5). Different uppercase letters indicate significant difference between different pathogenic fungis (p < 0.05). EF, endophyte-free A. sibiricum leaves; EG, Epichloë gansuensis-infected A. sibiricum leaves; ES, Epichloë sibiricum-infected A. sibiricum leaves; Bs, Bipolaris sorokiniana; Cl, Curvularia lunata; Ss, Sclerotinia sclerotioru; St, Sclerotinia trifoliorum.

表2 内生真菌(E)、病原真菌(P)和种植方式(M)对羽茅和羊草的病斑数以及病斑长度的多因素方差分析

Table 2 Multi-way ANOVA showing the effects of endophyte (E), pathogen (P) and mixture type (M) on lesion number and length of Achnatherum sibiricum and Leymus chinensis

因素 Factor	羽茅 Achnatherum sibiricum				羊草 Leymus chinensis
	病斑数 Lesion number		病斑长度 Lesion length		病斑数 Lesion number		病斑长度 Lesion length
		F	p	F	p	F	p	F	p
E	137.68	<0.001	65.73	<0.001
P	147.75	<0.001	3.16	0.078	82.59	<0.001	17.53	<0.001
M	1.32	0.271	1.83	0.165	4.87	0.011	0.81	0.451
P × M	0.02	0.980	0.55	0.578	1.24	0.298	0.20	0.819
E × P	4.74	0.032	2.33	0.130
E × M	0.13	0.871	0.31	0.734
E × P × M	0.26	0.774	0.15	0.861

图5 病原真菌侵染下不感染内生真菌(□)和感染内生真菌(■)羽茅的病斑数(A)和病斑长度(B)(平均值±标准误, n = 10)。不同小写字母表示差异显著(p < 0.05)。

Fig. 5 Mean number (A) and length of lesions (B) of endophyte-free (□) and endophyte-infected (■) Achnatherum sibiricum after inoculated by pathogenic fungi (means ± SE, n = 10). Different lowercase letters indicate significant difference (p < 0.05).

图6 病原真菌侵染下邻生植物羊草的病斑数(A)和病斑长度(B)(平均值±标准误, n = 10)。不同小写字母表示差异显著(p < 0.05)。Lc, 单种羊草; LcEF, 与EF为邻的混种羊草; LcEI, 与EI为邻的混种羊草。

Fig. 6 Mean number (A) and length of lesions (B) of neighbor Leymus chinensis inoculated by pathogenic fungi (means ± SE, n = 10). Different lowercase letters indicate significant difference (p < 0.05). Lc, L. chinensis grown in monoculture (mono); LcEF, L. chinensis grown with endophyte-free Achnatherum sibiricum; LcEI, L. chinensis grown with endophyte-infected A. sibiricum.

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