钙调节活性氧和内生细菌增强水稻对纹枯病抗性

doi:10.3864/j.issn.0578-1752.2025.11.006

摘要/Abstract

摘要：

【背景】由立枯丝核菌（Rhizoctonia solani）引起的水稻纹枯病在世界各稻区常年普遍发生，因抗性品种严重缺乏，给水稻绿色生态种植带来巨大挑战。【目的】揭示外源钙调物处理对水稻抗病的调节作用及其调控机制。【方法】四叶一心期籼稻和粳稻幼苗喷施5 mmol·L^-1钙调物150 mL，共喷施两次，每24 h喷施一次，在首次喷施48 h后采用牙签嵌入法在水稻第2、3叶片叶鞘接种立枯丝核菌，取同一部位叶片进行活性氧相关指标、光合色素含量测定及内生细菌分析。【结果】加钙（CaCl₂和Ca(NO₃)₂）处理增强供试水稻的抗病性，减钙（螯合剂EGTA）处理削弱水稻的抗病性。对各处理水稻植株钙含量分析发现，与空白对照相比，加钙处理提升植株钙含量，减钙处理降低植株钙含量；病原菌接种导致体内钙含量较相应对照处理均降低。加钙处理使水稻幼苗活性氧（和H₂O₂）含量增加，减钙处理引起的活性氧含量变化比加钙处理明显。加钙处理后，水稻活性氧含量变化与病斑面积变化趋势一致，二者正相关；SOD、POD、CAT、APX、GPX活性和GSH含量保持相对较高的水平，其中CAT活性增幅尤为明显。减钙处理多数情况下相关酶活性显著降低。加钙处理可重构水稻体内活性氧稳态，包括活性氧产生、抗氧化系统和膜损伤指标。与接种对照相比，加钙处理水稻病斑面积与H₂O₂含量正相关性增强，CAT负相关性增强、ASA负相关性增强、APX负相关性减弱、GSH由负相关转为正相关，MDA正相关性减弱、电导率由负相关转为正相关。加钙处理使接菌水稻叶片内生细菌群落微生物丰富度降低、内生细菌群落多样性有限增加。尽管加钙处理对水稻内生细菌共现网络影响有限，但立枯丝核菌接种对水稻内生细菌网络产生显著影响，且加钙处理内生菌网络鲁棒性在接菌前后均显著高于空白组和减钙组，接菌后网络易损性显著低于空白组和减钙组。【结论】加钙处理可以显著降低供试水稻相对病斑面积，提高水稻SOD、POD、CAT活性和GSH含量，减少体内活性氧积累，减少膜损伤，维持活性氧稳态提高水稻抗病性；同时，钙处理降低水稻内生细菌群落丰富度、增加其多样性，提高有益菌属占比，重构稳定性增强的内生细菌抗病群落。

关键词: 钙, 水稻, 纹枯病, 立枯丝核菌, 活性氧, 内生细菌

Abstract:

【Background】 Rice sheath blight, caused by Rhizoctonia solani, is a prevalent disease in rice-growing regions worldwide, posing significant challenges to the green ecological cultivation of rice due to the severe lack of resistant varieties.【Objective】 To elucidate the regulatory effect of exogenous calcium treatment on rice disease resistance and its regulatory mechanism.【Method】 Four-leaf stage indica and japonica rice seedlings were sprayed with a 5 mmol·L^-1 calcium regulator at 150 mL and sprayed twice at a 24 h interval. The toothpick insertion method was used to inoculate R. solani in the second- and third-leaf sheaths of rice 48 h after the first spray. Leaves from the same part were taken for determination of reactive oxygen species (ROS)-related indicators, photosynthetic pigment content, and endophytic bacterial analysis.【Result】 Calcium addition treatment (CaCl₂ and Ca(NO₃)₂) enhanced the resistance of the tested rice against R. solani while the chelating agent EGTA treatment weakened their resistance. Compared with the blank control, the calcium content of plants was increased with calcium addition treatment and decreased with calcium reduction treatment; pathogen inoculation resulted in lower calcium content in the plants compared to the corresponding control treatments. All calcium addition treatments caused an increase in the content of and H₂O₂ in rice seedlings, the change of ROS content caused by the calcium reduction treatment was more obvious than that of calcium addition treatment. Calcium addition treatment caused changes in rice ROS content consistent with the trend of lesion area changes, with a positive correlation between the two. Calcium addition treatment also maintained the enzyme activities of SOD, POD, CAT, APX, GPX and content of GSH at relatively high levels. The increase in CAT activity was particularly significant. In most cases, calcium reduction treatment significantly reduced the activity of related enzymes. Calcium addition treatment could reconstruct rice’s ROS homeostasis and enhance disease resistance and immune responses, including ROS production, antioxidant systems, and membrane damage indicators. Compared with the blank control, the positive correlation between the lesion area and H₂O₂ content in rice treated with calcium addition was enhanced, the negative correlation with CAT was enhanced, the negative correlation with ASA was enhanced, the negative correlation with APX was weakened, the correlation with GSH turned from the negative to positive, the positive correlation with MDA was weakened, and the correlation with conductivity turned from negative to positive. Calcium addition treatment reduced the microbial richness of endophytic bacterial community in inoculated rice leaves and limitedly increased the diversity of endophytic bacterial community. Although calcium addition treatment had a limited impact on the endophytic bacterial co-occurrence network in rice, the inoculation of R. solani significantly affected the endophytic bacterial network in rice. Furthermore, the network robustness after calcium addition treatment was significantly higher than that of the control group and the calcium reduction treatment group, both before and after inoculation; the network vulnerability after inoculation was significantly lower than that of the control group and the calcium reduction treatment group.【Conclusion】 Calcium addition treatment can significantly reduce the relative lesion area of tested rice, enhance the activities of SOD, POD, CAT and content of GSH, decrease the accumulation of ROS, reduce membrane damage, maintain the homeostasis of ROS and enhance the disease resistance of rice; simultaneously, calcium addition treatment reduces the richness of endophytic bacterial community, increases its diversity, enhances the proportion of beneficial bacterial genera, restructures the endophytic bacterial community with enhanced stability.

Key words: calcium, rice, sheath blight, Rhizoctonia solani, reactive oxygen species (ROS), endophytic bacteria

赵守帅, 都梦翔, 郭思宇, 张烁, 赵宏伟, 赵长江. 钙调节活性氧和内生细菌增强水稻对纹枯病抗性[J]. 中国农业科学, 2025, 58(11): 2145-2161.

ZHAO ShouShuai, DU MengXiang, GUO SiYu, ZHANG Shuo, ZHAO HongWei, ZHAO ChangJiang. Calcium Regulates Reactive Oxygen Species and Endophytic Bacteria to Enhance Rice Resistance to Sheath Blight[J]. Scientia Agricultura Sinica, 2025, 58(11): 2145-2161.

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引物（基因号） Primer (Gene ID)	引物序列 Primer sequence (5′-3′)
OsCAT2-F (LOC_Os06g51150)	GCACAGTTTGACAGGGAG
OsCAT2-R (LOC_Os06g51150)	GGTCTGAACACCAGGAGC
OsCNGC9-F (LOC_Os09g38580)	ACTCATGGTGGCCAGAAGAA
OsCNGC9-R (LOC_Os09g38580)	TGCATTTGAGCAATGGCACT
OsAnn8-F (LOC_Os09g20330)	CGATGTGCGACGCCAAGAC
OsAnn8-R (LOC_Os09g20330)	TCGCCGTCCTTCCTCTTCAG
OsMCA1-F (LOC_Os03g06120)	TTGATGTGTTGTTGCTGTGCG
OsMCA1-R (LOC_Os03g06120)	ACTGAAATGAGGGTGGGCTAATC
OsActin-F (LOC_Os10g36650)	CATGCTATCCCTCGTCTCGACCT
OsActin-R (LOC_Os10g36650)	GCACTTCATGATGGAGTTGTAT

处理 Treatment	相对病斑面积Relative lesion area
处理 Treatment	晶两优Jingliangyou	垦粳Kenjing
Con	0.32±0.01b	0.49±0.02b
CaCl₂	0.17±0.01d	0.08±0.01d
Ca(NO₃)₂	0.26±0.01c	0.16±0.01c
EGTA	0.41±0.01a	0.61±0.02a

测定指标 Measurement indicator	处理 Treatment	晶两优Jingliangyou		垦粳Kenjing
测定指标 Measurement indicator	处理 Treatment	BRS	ARS	BRS	ARS
含量 content (nmol·g^-1 FW)	Con	0.80±0.02c	1.55±0.02b**	0.22±0.01c	0.51±0.02b**
	CaCl₂	0.94±0.04b	1.21±0.01c**	0.27±0.01b	0.33±0.02d
	Ca(NO₃)₂	1.02±0.03b	1.26±0.01c**	0.29±0.01b	0.42±0.02c**
	EGTA	1.31±0.01a	1.67±0.06a**	0.43±0.03a	0.82±0.04a**
H₂O₂含量 H₂O₂ content (mmol·g^-1 FW)	Con	0.74±0.01b	1.25±0.02b**	0.58±0.02c	1.25±0.01b**
	CaCl₂	0.77±0.01b	0.88±0.01d**	0.66±0.02b	0.86±0.02d**
	Ca(NO₃)₂	0.77±0.01b	0.96±0.01c**	0.70±0.01b	0.94±0.02c**
	EGTA	0.86±0.02a	1.61±0.02a**	1.04±0.03a	1.43±0.02a**
电解质渗透率 Electrolyte leakage rate (%)	Con	6.20±0.18b	12.15±0.18a**	3.88±0.19b	5.68±0.24b**
	CaCl₂	5.40±0.10c	7.61±0.12c**	3.02±0.16c	3.66±0.13d
	Ca(NO₃)₂	5.28±0.15c	10.18±0.11b**	3.52±0.19bc	4.41±0.20c
	EGTA	7.99±0.22a	11.69±0.15a**	4.57±0.19a	8.60±0.22a**
MDA含量 MDA content (µmol·g^-1 FW)	Con	0.0091±0.0002b	0.0137±0.0001b**	0.0060±0.0001b	0.0090±0.0001b**
	CaCl₂	0.0089±0.0001b	0.0119±0.0002c**	0.0047±0.0001c	0.0056±0.0002d**
	Ca(NO₃)₂	0.0093±0.0002b	0.0118±0.0002c**	0.0060±0.0001b	0.0069±0.0002c**
	EGTA	0.0121±0.0002a	0.0161±0.0001a**	0.0091±0.0001a	0.0113±0.0002a**

测定指标 Measurement indicator	处理 Treatment	晶两优Jingliangyou		垦粳Kenjing
测定指标 Measurement indicator	处理 Treatment	BRS	ARS	BRS	ARS
Ca含量 Ca content (μg·g^-1)	Con	218.51±7.93b	113.55±5.43b**	92.89±6.35c	66.99±3.60b
	CaCl₂	242.76±8.96a	143.31±10.88a**	102.53±10.66b	86.28±2.63a
	Ca(NO₃)₂	233.67±9.17a	136.14±8.02ab**	124.30±6.04a	72.50±4.54b**
	EGTA	189.31±8.84c	160.11±7.90a	86.00±7.57d	71.68±4.51b
SOD活性 SOD activity (U·g^-1 FW)	Con	108.80±1.37c	142.69±1.52c**	140.37±0.93b	166.67±1.26b**
	CaCl₂	146.07±0.90a	172.83±0.91a**	140.89±0.55b	174.28±1.70a**
	Ca(NO₃)₂	149.17±1.02a	169.86±0.26a**	146.19±1.54a	174.02±0.68a**
	EGTA	125.88±1.92b	163.17±1.10b**	136.29±0.63c	131.60±1.99c
POD活性 POD activity (U·mg^-1 FW)	Con	24.58±0.18c	26.94±0.29b**	26.46±0.26c	30.13±0.48c**
	CaCl₂	27.91±0.36b	30.76±0.41a**	31.29±0.13a	34.57±0.30a**
	Ca(NO₃)₂	29.10±0.20a	30.16±0.64a	28.50±0.03b	32.07±0.28b**
	EGTA	22.98±0.20d	26.38±0.44b**	25.67±0.07d	30.88±0.15c**
CAT活性 CAT activity (U·g^-1 FW)	Con	28.89±2.22b	46.67±3.33b	28.89±2.22b	71.11±2.22b**
	CaCl₂	42.22±2.22a	93.33±3.85a**	55.56±2.22a	100.00±3.85a**
	Ca(NO₃)₂	37.78±2.22a	80.00±3.85a**	55.56±2.22a	95.56±2.22a**
	EGTA	22.22±2.22b	33.33±6.67b	22.22±2.22b	68.89±2.22b**

测定指标 Measurement indicator	处理 Treatment	晶两优Jingliangyou		垦粳Kenjing
测定指标 Measurement indicator	处理 Treatment	BRS	ARS	BRS	ARS
APX活性 APX activity (U·g^-1 FW)	Con	70.00±3.21b	84.22±2.25a**	81.48±1.48b	106.67±2.57c**
	CaCl₂	78.89±1.92a	88.07±1.78a	79.26±3.70b	144.44±4.44a**
	Ca(NO₃)₂	83.33±1.92a	89.56±1.90a	97.78±6.67a	133.33±2.57b**
	EGTA	55.56±1.28c	42.22±1.28b**	46.67±2.57c	76.30±2.67d**
GPX活性 GPX activity (U·mg^-1 FW)	Con	1.72±0.02b	2.20±0.01b**	2.27±0.01c	2.34±0.02b**
	CaCl₂	1.93±0.02a	2.38±0.002a**	2.32±0.005b	2.43±0.01a**
	Ca(NO₃)₂	1.91±0.01a	2.15±0.02c**	2.36±0.02a	2.42±0.02a
	EGTA	1.52±0.005c	2.06±0.02d**	2.24±0.01c	2.29±0.01c
ASA含量 ASA content (μg·g^-1 FW)	Con	208.55±3.03a	195.83±4.91a	207.81±0.52a	207.18±0.61a
	CaCl₂	208.63±3.79a	196.35±3.08a	208.90±0.52a	207.41±0.69a
	Ca(NO₃)₂	209.28±2.61a	196.19±3.67a	209.33±0.6a	208.88±0.58a
	EGTA	187.11±3.67b	172.46±1.77b	206.08±0.45b	203.15±0.63b
GSH含量 GSH content (μmol·g^-1 FW)	Con	308.14±4.14c	320.73±4.64c	349.52±3.18ab	370.06±3.60b
	CaCl₂	330.67±4.03b	373.08±5.78b**	360.12±3.44a	383.98±5.74b
	Ca(NO₃)₂	351.21±5.78a	406.21±4.64a**	360.79±5.18a	419.10±10.35a**
	EGTA	333.32±1.75b	296.87±2.89d**	338.68±3.44b	321.27±3.60c