松材线虫生防真菌伊氏线虫菌研究进展

doi:10.11707/j.1001-7488.20200118

摘要/Abstract

摘要：

松材线虫病对森林生态健康有显著影响，可以导致松林大面积死亡，有效预防松材线虫病是维护森林生态系统的重要措施。伊氏线虫菌是一种松材线虫的内寄生真菌，具有作为松材线虫病生物防治的天敌的潜力。近几年，对于这种鲜为人知的真菌的研究增多。本文从该菌的发现、形态学、培养、侵染机制、分子层面研究和生物防治应用等方面较系统全面地介绍该菌。伊氏线虫菌共有6个菌株，能产生月形孢子和杆状孢子2种类型的孢子，月形孢子能黏附到松材线虫体表并寄生于体内将其杀死。伊氏线虫菌能在松树和松树分泌的树脂中存活，通过产生宿主松树的香味而引诱松材线虫。培养条件对伊氏线虫菌生长、产孢、孢子侵染活力的影响较大，可以在培养基中添加甘氨酸等增强其对环境的抗性。伊氏线虫菌基因组已被完整测序，还克隆出1个对松材线虫具有强毒力的丝氨酸蛋白酶基因，同时在菌株细胞内发现内生细菌的存在，可以通过分子方法检测该菌。在温室和野外采用树干注射、树冠喷洒和伤口接种等不同方法施用含有该真菌的生防菌，均显示其能降低松材线虫对松树的侵染。目前对于该菌的研究主要集中在实验室，对于林间防治应用也有研究，但还是较少。关于该菌未来的研究可以从几方面开展：1）分子层面弄清其侵染机制；2）商品化生物制剂；3）野外防治方案；4）松褐天牛与伊氏线虫菌之间的关系。

关键词: 伊氏线虫菌, 松材线虫, 生物防治

Abstract:

Pine wilt disease has a significant impact on the forest ecological health, which can lead to large-scale death of pine forests. Effective prevention of pine wood nematode disease is an important measure to maintain the forest ecosystem. Esteya vermicola is an endoparasitic fungus of Bursaphelenchus xylophilus, a pine wood nematode, and has potential as a biological control agent for pine wood nematode disease. In recent years, the research on this little-known fungal species has increased. Based on this, this article systematically introduces the bacteria from the aspects of its discovery, morphology, culture, infection mechanism, molecular level research and biological control applications. E. vermicola, a total of six strains, can produce two types of spores:lunate spores and rod-shaped spores. Moon spores can adhere to the surface of pine wood nematode and parasitize the nematode to kill nematodes. E. vermicola can survive in pine and in the resin secreted by pine, and attracts pine wood nematodes by producing the scent of host pine trees. The culture conditions have a great influence on the growth, sporulation, and spore invasion of E. vermicola. Glycine can be added to the medium to enhance its resistance to the environment. The genome of E. vermicola has been completely sequenced, and a serine protease gene with highly virulent to B. xylophilus has been cloned. Endophytic bacteria are also found in the cells of the strain, which can be detected by molecular methods. The application of biocontrol containing the fungi in different ways, such as trunk injection, canopy spraying and wound inoculation in the greenhouse and in the field, showed that it can reduce the infection of the pine wood nematode to pine wood. At present, the research on this fungus is mainly concentrated in the laboratory. There are also studies on the application of field prevention and control, but it is still relatively less. The research of this fungus in the future could be carried out in several aspects:1) molecular level research to clarify its infestation mechanism; 2) research on the commercial biological agent production of the fungus; 3) research on field prevention and control programs; 4) the research on the relationship between Monochamus alternatus and E. vermicola.

Key words: Esteya vermicola, Bursaphelenchus xylophilus, biological control

中图分类号:

S718.81

茅裕婷,马涛,蓝来娇,温秀军. 松材线虫生防真菌伊氏线虫菌研究进展[J]. 林业科学, 2020, 56(1): 180-190.

Yuting Mao,Tao Ma,Laijiao Lan,Xiujun Wen. Advances in Esteya vermicola, A Potential Biocontrol Fungus for Pine Wood Nematode[J]. Scientia Silvae Sinicae, 2020, 56(1): 180-190.

图/表 3

表1

表2

4个E. vermicola菌株的形态特征比较"

项目Item	ATCC74485 (Liou et al., 1999)	CBS115803 (Kubátová et al.，2000)	CNU120806 (Wang et al.，2008)	NKF13222 (王婷婷，2014b)
PDA上生长速度(1周)Growth rate on PDA (one week)	直径3.0~4.5 cm，生长速度适中3.0-4.5 cm in diameter, moderate growth rate.	直径3.0~3.7 cm，生长速度较慢。3.0-3.7 cm in diameter, slow growth.	直径4.0~4.5 cm，生长速度较快。4.0-4.5 cm in diameter, fast growth.	直径3.1~3.5 cm，生长速度较慢。3.1-3.5 cm in diameter, slow growth.
PDA上菌落颜色Colony color on PDA	正面灰色、灰绿色至暗绿色，反面绿色至深绿色。Positive gray, grayish green to dark green, reverse green to dark green.	正面灰色至灰绿色，反面深绿色。Positive gray to grayish green, reverse dark green.	正反都为深绿色。Both positive and negative are dark green.	正面灰色、灰绿到黑绿色，反面灰色至深灰色。Positive gray, grayish green to dark green, reverse gray to dark gray.
产孢种类Spore-forming species	月形孢子：1个分生孢子梗产生1个孢子，(8.2~11.1)μm×(3.5~3.7)μm;杆状孢子：(4.4~7.4)μm×(1.5~1.9)μm。Lunar spores: One conidial stem produces one spore, (8.2-11.1) μm × (3.5-3.7) μm; Rod-shaped spores: (4.4-7.4) μm × (1.5-1.9) μm.	月形孢子：1个分生孢子梗产生1~4个孢子，(9.3~12.4)μm×(3.0~3.2)μm;杆状孢子：(4.6~6.2)μm×(1.3~1.5)μm。Lunar spores: one conidial stem produces 1~4 spores, (9.3-12.4) μm × (3.0-3.2) μm; Rod-shaped spores: (4.6-6.2) μm × (1.3-1.5) μm.	月形孢子：1个分生孢子梗产生1~4个孢子，(7.7~12.1)μm×(3.0~3.8)μm;杆状孢子：(4.3~8.4)μm×(3.0~3.8)μm。Lunar spores: one conidial stem produces 1-4 spores, (7.7-12.1) μm × (3.0-3.8) μm; Rod-shaped spores: (4.3-8.4) μm × (3.0-3.8) μm.	月形孢子：1个分生孢子梗产生一个孢子，(8.7~11.9)μm×(3.0~3.6)μm;杆状孢子：(4.3~7.5)μm×(1.3~2.1)μm。Lunar spores: One conidial stem produces one spore, (8.7-11.9) μm × (3.0-3.6) μm; Rod-shaped spores: (4.3-7.5) μm × (1.3-2.1) μm.
月形孢子萌发方式Germination method of moon spore	从孢子凸面萌发并产生一个芽管。Germination from spores convex and generating a germ tube.	从孢子凹面中部萌发并产生一个或多个芽管。Germinates from the middle of the concave surface of the spore and produces one or more germ tubes.	主要从孢子凹面萌发，偶尔可见凸面萌发。Mainly germinated from the concave surface of spores, occasionally visible convex germination.	从孢子凹面中部萌发并产生一个芽管。Germinates and produces a germ tube from the middle of the concave surface of the spore.
杆状孢子萌发方式Germination method of rod-shaped spores	从孢子端部萌发，也可从孢子1/3~1/2处萌发，可产生1到多个芽管。Germination from the ends of spores can also germinate from 1/3-1/2 spores, producing one or more germ tubes.	从孢子端部萌发产生多个芽管。Germination of multiple germ tubes from the ends of spores	90%从孢子1/3~1/2处萌发，10%从孢子端部萌发。90% germinated from 1/3-1/2 spores and 10% germinated from spore tips	从孢子端部萌发，也可从孢子1/3~1/2处萌发，可产生一到多个芽管。Germination from the ends of spores can also germinate from 1/3-1/2 spores, producing one or more germ tubes
营养条件对产孢的影响。BR Effect of nutritional conditions on sporulation	在营养丰富的培养基中主要产生杆状孢子，营养匮乏的培养基中主要产生新月形孢子。Produces rod-shaped spores in nutrient-rich cultures, producing crescent-shaped spores in nutrient-poor media.	在不同营养条件的培养基中2种孢子均产生较早。Both spores produced earlier in culture medium with different nutrient conditions	在营养丰富的培养基中主要产生新月形孢子。Produces crescent-shaped spores in a nutrient-rich medium	在营养丰富的培养基中主要产生杆状孢子，营养匮乏的培养基中主要产生新月形孢子。Produces rod-shaped spores in nutrient-rich cultures, producing crescent-shaped spores in nutrient-poor media

表2

表3

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菌株号 Strain No.	分离基质 Separation Matrix	来源 Source	分离年份 Year	参考文献References
ATCC74485	被松材线虫侵染的黑松心材木片 Black pine wood chips infested by B. xylophilus	中国台湾 Taiwan, China	1995	Liou et al., 1999
CBS115803	正毛束小蠹幼虫、成虫和其在橡树的坑道 Larvae, adult beetles of S. intricatus and their galleries in oak	捷克 Czech Republic	1999	Kubátová et al., 2000
CNU120806	土壤中被侵染的腐生线虫 Saprophytic nematodes infested by E. vermicola in the soil	韩国 Korea	2006	Wang et al., 2008
NKF132229	包装木中被侵染的鲁尔夫伞滑刃线虫 B. rainulgi infected by E. vermicola in packaging wood	巴西 Brazil	2014	王婷婷，2014b
CBS156.82	干燥的松属植物Dry Pinus	日本Japan	1982	Wang et al., 2009a
CBS100821	油橄榄嫩枝Olive twig	意大利Italy	1998	Wang et al., 2009a

线虫Nematodes	分生孢子Lunate conidia		参考文献References
线虫Nematodes	黏附Attachment	侵染Infection	参考文献References
水稻干尖线虫Aphelenchoides besseyi	+	+	曾显雄，2003; Zeng, 2003
叶芽线虫Aphelenchoides sp.	+	+	曾显雄，2003; Zeng, 2003
松材线虫B. xylophilus	+	+	曾显雄，2003; Zeng, 2003
拟松材线虫B. mucronatus	+	+	王婷婷，2014b; Wang, 2014b
腐烂茎线虫Ditylenchus destructor	+	+	王婷婷，2014b; Wang, 2014b
稻茎线虫D. angustus	+	+	Tzean et al.，2001
甜菜茎虫D. dipsacis	+	+	Tzean et al.，2001
椰子红环线虫Phadinaphelenchus cocophilus	+	+	Tzean et al.，2001
草莓滑刃线虫A. fragariae	+	+	Tzean et al.，2001
全齿复活线虫Panangrellus redivivus	+	+	Wang et al.，2008
肾形线虫Rotylenchulus reniformis	+	-	曾显雄，2003; Zeng, 2003
根腐线虫Pratylenchus coffeae	-	-	曾显雄，2003; Zeng, 2003
南方根瘤线虫Meloidigyne incognita	-	-	曾显雄，2003; Zeng, 2003
燕麦孢囊线虫Heterodera avenae	-	-	王婷婷，2014b; Wang, 2014b
穿刺短体线虫Pratylenchus penetrans	-	-	王婷婷，2014b; Wang, 2014b

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