干旱胁迫及外生菌根菌对马尾松幼苗根系形态及分泌物的影响

doi:10.11707/j.1001-7488.20220707

摘要/Abstract

摘要：

目的: 探究干旱胁迫下接种不同外生菌根真菌对马尾松幼苗根系生长、形态、根系分泌物及根际土壤酶活性的影响，为阐明不同外生菌根菌促生抗逆的策略提供参考。方法: 采用棚内盆栽试验，分析正常浇水和干旱胁迫条件下，接种橙黄硬皮马勃(Sc)、琥珀乳牛肝菌(Sp)和未接菌(CK)马尾松幼苗的地上和地下生长、根系分泌物及根际土壤酶活性差异。结果: 1) 干旱胁迫抑制马尾松生长和根系活力，但接种Sc和Sp均能促进马尾松幼苗生长、生物量积累和根系活力，改善根系形态，以接种Sc效果更佳。2) 在根系分泌物中共检测出酸类、酮类、萜类、酯类、醇类和醛类6大类有机物质，共28种化合物。分泌物组成和相对含量受接菌和干旱胁迫调控，其中，接种Sc和Sp分别使分泌物种类增加和降低；无论是否接菌，分泌物种类均在干旱胁迫下增加；各类分泌物相对含量对干旱和接菌处理响应明显且存在差异，变化幅度和方向与分泌物种类有关；接种菌根菌能改变干旱胁迫下各类分泌物相对含量的变化，主要表现在抑制萜类相对含量、增加醛类相对含量，同时，酸类物质相对含量下降程度更大。菌根菌对同一组分不同化合物相对含量也有显著影响。3) 马尾松幼苗接种Sp后通过提高根际土壤脲酶和蔗糖酶促进植物生长，抵御干旱胁迫；接种Sc后通过维持较高的酸性磷酸酶和脲酶活性抵御干旱胁迫。结论: 马尾松幼苗接种Sp和Sc通过改善根系形态、提高根系活力和维持较高的土壤酶活性等方式促进生长和缓解干旱胁迫，以接种Sc缓解干旱胁迫的效果最佳。不同菌根菌接种处理后根系性状及其在干旱胁迫下的变化复杂且存在差异。

关键词: 马尾松, 外生菌根, 干旱, 根系分泌物, 土壤酶

Abstract:

Objective: To study the effects of different ectomycorrhizal fungi inoculation under drought stress on the root growth, root exudates and rhizosphere soil enzyme activities of Pinus massoniana seedlings, and to understand the strategies of different ectomycorrhizal fungi to promote growth and resist stress. Method: With the pot experiment in the greenhouse and two conditions of normal watering and drought stress, the difference in growth of above and below ground, root exudates and rhizosphere soil enzyme activities of different Pinus massoniana seedlings that respectively inoculated of Suillus placidus (Sp), Scleroderma citrinum (Sc) and uninoculated (CK) were studied. Result: 1) Drought stress inhibited growth and root vitality of P. massoniana seedlings, but Sc and Sp inoculation promoted seedling growth, biomass accumulation and root vitality, and also improved root morphology, with better performances in Sc inoculation seedlings. 2) A total of 6 types of organic substances, including acids, ketones, terpenoids, esters, alcohols and aldehydes, were detected in the root exudates, with a total of 28 compounds. The composition and relative content of secretions were regulated by inoculation and drought stress. Sc inoculation can increased the types of exudates while Sp inoculation decreased that; Drought stress increased the types of exudates and was not affected by inoculation; The relative content of various exudates responds obviously and differently to drought and inoculation treatments, and the change trends were related to the types of exudates. At the same time, the two ectomycorrhizal fungi also changed the relative content of various secretions under drought stress, mainly in inhibiting of the relative content of terpenoids, increasing of the relative content of aldehydes, and decreasing more in the relative content of acids. The effects of the two ectomycorrhizal fungi on the relative content of different compounds of the same component were also significantly different. 3) Sp inoculation treatment promoted plant growth and resists drought stress by increasing rhizosphere soil urease and invertase, while Sc treatment resisted drought stress by maintaining higher acid phosphatase and urease activities. Conclusion: Inoculation of Sp and Sc promoted the growth of P. massoniana and alleviate drought stress by improving root morphology, increasing root vitality and maintaining higher soil enzyme activities. Sc inoculation had the best effect on alleviating drought stress. Root traits and their changes under drought stress were complex and different after different mycorrhizal fungi inoculation. This is of great significance for understanding the mechanism of mycorrhizal fungi regulating root exudates in response to drought stress.

Key words: Pinus massoniana, ectomycorrhiza, drought, root exudates, soil enzyme

中图分类号:

S718.43

李敏, 赵熙州, 王好运, 卢中科, 丁贵杰. 干旱胁迫及外生菌根菌对马尾松幼苗根系形态及分泌物的影响[J]. 林业科学, 2022, 58(7): 63-72.

Min Li, Xizhou Zhao, Haoyun Wang, Zhongke Lu, Guijie Ding. Effects of Drought Stress and Ectomycorrhizal Fungi on the Root Morphology and Exudates of Pinus massoniana Seedlings[J]. Scientia Silvae Sinicae, 2022, 58(7): 63-72.

图/表 7

图1

表1

表2

图2

表3

表4

不同处理中根系分泌物化学组分相对含量"

类别 Category	物质名称 Name of substances	分子式 Molecular formula	分子量 Molecular weight	相对含量Relative content (%)
				CK		Sc		Sp
				浇水 Watering	干旱 Drought	浇水 Watering	干旱 Drought	浇水 Watering	干旱 Drought
萜类Terpenoids	柠檬烯Limonene	C₁₀H₁₆	136	0.32	0.119	0.095	0	0	0
	β-水芹烯β-Phellandrene	C₁₀H₁₆	136	0.64	0.45	0.34	0.45	0.58	0.62
	长叶烯Longifolene	C₁₅H₂₄	204	2.69	3.65	1.71	0.95	5.22	1.55
酯类Esters	丙烯酸-2-乙基己酯 2-Ethylhexyl acrylate	C₁₁H₂₀O₂	184	0.62	0.22	0.94	0.73	0	0
酯类Esters	癸酸甲酯Methyl caprate	C₁₁H₂₂O₂	186	0.69	0.32	0.21	0.33	0	0.56
醛类Aldehyde	安息香醛Benzaldehyde	C₇H₆O	106	1.74	0.93	0.62	0.70	0.20	0.88
	5-甲基糠醛 5-methyl-2- furancarboxaldehyde	C₆H₆O₂	110	0.23	0.15	0.07	0.10	0.15	0.22
	3, 5-二甲基苯甲醛 3, 5-dimethyl-benzaldehyde	C₉H₁₀O	134	0.79	0.39	0.79	0.56	0.48	0.40
	香草醛Vanillin	C₈H₈O₃	152	0.50	1.00	0.43	0.61	0.59	1.61
酸类Acids	丙酸Propanoic acid	C₃H₆O₂	74	1.70	1.14	0.20	0.31	0.03	0.19
	2- 甲基丙酸 2-methyl-propanoic acid	C₄H₈O₂	88	0.36	0.57	0.06	0.13	0	0
	丁酸Butanoic acid	C₄H₈O₂	88	0.56	0.09	0.76	0.83	0.30	0.28
	3-甲基丁酸 3-Methyl-butanoic acid	C₅H₁₀O₂	102	0.82	1.18	0.59	0.32	0	0.47
	丁烯酸Isocrotonic acid	C₄H₆O₂	86	0.76	1.91	0.72	1.65	0.68	1.87
	辛酸Octanoic acid	C₈H₁₆O₂	144	0	0.29	0	0	0.59	0
	壬酸Nonanoic acid	C₉H₁₈O₂	158	0.19	0.28	0.37	0.19	0.50	0.40
	(E)-2-辛烯酸 (E)-2-octenoic acid	C₈H₁₄O₂	142	0	0	0.11	0.12	0	0
	正癸酸n-Decanoic acid	C₁₀H₂₀O₂	172	0	0	0	0	0.47	0
	苯甲酸Benzoic acid	C₇H₆O₂	122	0.43	0.50	0.49	0.55	0.76	0.72
	月桂酸Dodecanoic acid	C₁₂H₂₄O₂	200	0.78	0.67	0.47	0.77	1.24	0.81
	苯乙酸Benzeneacetic acid	C₈H₈O₂	136	1.35	5.79	1.27	4.13	2.04	5.99
	十四烷酸Tetradecanoic acid	C₁₄H₂₈O₂	228	1.30	2.05	2.97	3.30	9.25	12.30
	十五烷酸Pentadecanoic acid	C₁₅H₃₀O₂	242	1.11	1.20	2.63	2.17	3.16	4.65
	十六烷酸n-hexadecanoic acid	C₁₆H₃₂O₂	256	49.82	40.75	45.98	40.04	36.85	26.66
	硬脂酸Octadecanoic acid	C₁₈H₃₆O₂	284	25.40	23.70	26.16	24.30	20.12	14.92
	油酸Oleic acid	C₁₈H₃₄O₂	282	1.97	5.25	5.52	6.19	10.07	8.40
醇类Alcohol	苯甲醇Benzyl alcohol	C₇H₈O	108	0.85	0.71	0.60	0.65	0	0
酮类Ketones	六氢法尼基丙酮 Hexahydrofarnesyl acetone	C₁₈H₃₆O	268	0.47	0.56	0.20	0.57	0.78	1.13

表4

图3

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处理 Treatments	苗高净增长量 Growth of height/cm	地径净增长量 Growth of stem diameter/mm	根生物量 Dry weight of root/(g·plant^-1)	地上生物量 Dry biomass of aboveground/ (g·plant^-1)	总生物量 Dry biomass/(g·plant^-1)	根冠比 Root aboveground ratios	侵染率 Infection rate (%)
CKW	5.13±0.18b	1.54±0.02b	0.7±0.05Ab	4.24±0.43Ab	4.95±0.94Ab	0.17±0.02	3.17±0.65e
CKD	1.93±0.07d	0.5±0.01d	0.33±0.04Bb	1.67±0.14Bb	2.00±0.38Bb	0.20±0.02	1.07±0.12e
SpW	5.63±0.14a	1.65±0.01a	1.26±0.12Aa	5.57±0.66Aa	6.83±1.54Aa	0.23±0.01	63.98±1.72b
SpD	2.13±0.09d	0.53±0.02d	0.57±0.08Bab	2.57±0.60Bab	3.15±0.63Bab	0.25±0.08	41.39±1.70d
ScW	5.67±0.24a	1.67±0.01a	1.49±0.21Aa	6.50±0.98Aa	7.99±1.07Aa	0.24±0.04	68.52±1.60a
ScD	3.17±0.39c	0.64±0.02c	0.72±0.13Ba	3.10±0.42Ba	3.82±0.44Ba	0.25±0.06	49.26±0.91c
接菌ECM	< 0.001	< 0.001	< 0.001	< 0.001	< 0.001	0.228	< 0.001
干旱Drought(D)	< 0.001	< 0.001	< 0.001	< 0.001	< 0.001	0.56	< 0.001
ECM×D	< 0.001	0.041	0.232	0.744	0.579	0.97	< 0.001

处理 Treatments	根长 Length/cm	根表面积 Surfale area/cm²	平均直径 Avgrage diam/mm	根体积 Root volume/cm³	根尖数 Tips	分支数 Forks
CKW	697.92±12.54b	105.23±4.87b	2.23±0.08d	1.60±0.06c	6 626.75±466.16c	12 614.00±353.02c
CKD	232.86±18.60cd	56.80±3.23c	4.23±0.34c	0.81±0.08d	2 168.33±205.85e	6 643.33±367.73c
SpW	1454.13±78.76a	252.68±16.82a	7.36±0.61b	3.62±0.17b	14 155.33±485.14b	36 551.00±1 031.20b
SpD	248.91±30.44d	52.65±7.45c	1.22±0.11e	0.90±0.07d	5 789.33±283.58cd	10 349.00±610.07c
ScW	1456.40±116.87a	251.00±15.92a	7.77±0.28b	5.16±0.21a	27 409.33±2 303.59a	73 145.33±4 766.79a
ScD	490.67±17.46c	89.57±2.18b	10.06±0.29a	1.69±0.14c	2 991.67±234.52e	8 395.67±327.10c
接菌ECM	< 0.001	< 0.001	0.033	< 0.001	< 0.001	< 0.001
干旱Drought(D)	< 0.001	< 0.001	< 0.001	< 0.001	< 0.001	< 0.001
ECM×D	< 0.001	< 0.001	< 0.001	< 0.001	< 0.001	< 0.001

类别 Category	种类数Number of kind
	CK		Sc		Sp
	浇水Watering	干旱Drought	浇水Watering	干旱Drought	浇水Watering	干旱Drought
萜类Terpenoids	3.65(3)	4.22(3)	2.14(3)	1.40(2)	5.80(2)	2.16(2)
酯类Esters	1.32(2)	0.54(2)	1.15(2)	1.06(2)	0	0.56(1)
醛类Aldehyde	3.25(4)	2.47(4)	1.91(4)	1.97(4)	1.42(4)	3.11(4)
酸类Acids	86.55(14)	85.37(15)	88.26(15)	84.99(15)	86.04(14)	77.67(13)
酮类Ketones	0.47(1)	0.56(1)	0.20(1)	0.57(1)	0.78(1)	1.13(1)
醇类Alcohol	0.85(1)	0.71(1)	0.60(1)	0.65(1)	0	0
其他Other	3.91(3)	6.12(4)	5.73(4)	9.35(6)	5.96(4)	15.37(6)
小计(种)Total(Species)	28	30	30	31	25	27

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