水稻类病变突变体性状及分子机理研究进展

doi:10.16819/j.1001-7216.2018.7153

摘要/Abstract

摘要：

类病变突变体是植物在无显著逆境或损伤情况下以及未受到病原物危害时叶片上自然形成的类似病斑的一类突变体。水稻类病变突变体的产生常伴有抗病性的增强,其中涉及到防御相关基因的表达。相关研究主要集中于各种突变体的定位和克隆以及突变体的抗病性和抗病机制。综述了水稻类病变突变基因的性状及抗病性的研究进展,特别是对水稻类病变突变体形成原因、遗传特性、调控途径进行了梳理,并提出对水稻抗病育种的展望,以期为进一步分析类病变突变体的各种机制奠定理论基础,并为水稻育种提供参考。

关键词: 水稻类病变, 性状, 机理, 进展

Abstract:

Lesion mimic mutants are similar withnaturally-formed scab on rice leaveswithout significant stress or damage and without exposure to pathogens. The cause of lesion mimic traits in rice is often accompanied by increased resistance to disease, which involves the expression of defense-related genes. The related research mainly focuses on the localization and cloning of various mutants, as well as the disease resistance and its mechanism. We reviewed the research progress in lesion mimic mutant gene traits and disease resistance in rice, especially explaining the causes, genetic characteristics and regulatory pathways of lesion mimic mutants in rice,and expecting to improve rice breeding for disease resistance as well. All these will lay a theoretical foundation for further analysis of various mechanisms of lesion mimic mutants and providing some reference for rice breeding.

Key words: rice lesion mimic, traits, mechanisms, progress

中图分类号:

焦然, 徐娜, 胡娟, 宋周琳, 胡佳青, 饶玉春, 王跃星. 水稻类病变突变体性状及分子机理研究进展[J]. 中国水稻科学, 2018, 32(3): 285-295.

Ran JIAO, Na XU, Juan HU, Zhoulin SONG, Jiaqing HU, Yuchun RAO, Yuexing WANG. Advances in Traits of Lesion Mimic Mutants and Its Molecular Mechanisms in Rice[J]. Chinese Journal OF Rice Science, 2018, 32(3): 285-295.

图/表 4

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突变体 Mutant	病斑表型特征 Phenotype of lesion	突变体来源 Origin	染色体 Chr.	遗传方式^a	文献 Ref.
spl1	播种后15~20d,叶片或者叶鞘出现橙黄色类病斑并扩散到整个叶片	Spontaneous	12	SR	[6]
spl2	苗期叶片和茎表面出现不明显的变色,分蘖期变色更加明显	Spontaneous	2	SR	[11]
spl3	在分蘖期到抽穗期,叶片上出现较小的红色类病斑	γ-ray	3	SR	[11]
spl4	叶片上零散出现红棕色类病斑	γ-ray	6	SR	[11]
spl5	在分蘖期到抽穗期,叶片上出现较小的红色类病斑	γ-ray	7	SR	[12]
spl6	分蘖期叶片上出现较大的棕色类病斑	MNU	1	SR	[13]
spl7	在分蘖期到抽穗期,叶片上出现较小的红色类病斑	γ-ray	5	SR	[14]
spl8	分蘖期后,整个叶片表面都会出现红棕色类病斑	MNU	5	SR	[11]
spl9	植株花期后,在叶片和茎上会有较小的黑棕色类病斑	γ-ray	7	SR	[11]
spl10	在叶片上有黄色或棕色的类病斑	MNU	10	SR	[11]
spl11	分蘖期后出现铁锈样的类病斑,然后到成熟期后,病斑蔓延到整个叶片	EMS	12	SR	[15]
spl12	苗期,叶片枯萎且表现出黄绿色	MNU	-	SD	[16]
spl13	播种后10周后,植株叶片出现几毫米长的棕色类病斑	MNU	-	SD	[16]
spl14	播种后10周,叶片上出现暗棕色类病斑	MNU	-	SR	[16]
spl15	播种后3~4周,出现5~10mm的淡棕色类病斑	MNU	-	SD	[16]
spl16	叶片上出现小的黑色类病斑	DEB	-	SR	[10]
spl17	叶片上出现小的棕褐色类病斑	DEB	-	SR	[10]
spl18	叶片上出现小的棕褐色类病斑	T-DNA	10	SD	[10]
spl19	叶片上出现棕黄色类病斑,叶片变卷曲,变窄	DEB	-	SR	[10]
spl20	叶片上出现白色类病斑	FN	-	SR	[10]
spl21	叶片上出现棕黄色类病斑	DEB	12	SR	[10]
spl22	叶片上出现黑色类病斑	DEB	-	SR	[10]
spl23	叶片上出现黄色类病斑	DEB	-	SR	[10]
spl24	叶片上出现多而密集的黑色斑点	DEB	-	SD	[10]
spl25	叶片上出现深绿色斑点	DEB	-	SR	[10]
spl26	基因纯合致死,叶片出现红褐色类病斑	DEB	-	SD	[10]
spl27	基因纯合致死,叶片出现棕黄色类病斑	DEB	-	SD	[10]
spl28	植株产生类病斑的同时伴随植株的严重早衰,病斑呈黄褐色	MNU	1	SR	[17]
spl30	叶片出现红褐色类病斑并逐渐扩散	EMS	3	SR	[18]
spl30(t)	叶片出现红棕色类病斑	Transgene	12	SR	[18]
spl31	叶片出现棕色类病斑	EMS	12	SR	[19]
spl32	叶片边缘出现棕黄色类病斑	-	11	SD	[20]
bl1	叶片上首先出现棕色病变斑点,然后蔓延到穗上	Spontaneous	1	SR	[11]
bl2	叶片出现黑色类病斑并逐渐蔓延扩散	Spontaneous	2	SR	[11]
bl3	叶片上出现黑色类病斑,植株成熟后类病斑会蔓延到颖壳	Spontaneous	6	SR	[11]
bl4	植株上大范围出现大的棕色类病斑	32P	3	SR	[11]
bl5	叶片上出现很小的点状棕色类病斑	32P/X-ray	-	SR	[11]
bl6	叶片上出现较大的棕色类病斑	32P	-	SR	[11]
cdr1	叶片上出现类病斑,为起始型	MNU	-	SR	[21]
cdr2	起始型类病斑	MNU	-	SR	[21]
cdr3	基因纯合致死	MNU	-	SD	[21]
Fgl	叶片伸长过程中,叶片出现黄色或者白色的类病斑	Spontaneous	10	SR	[22]
Zn	叶片上出现黄色类病斑	γ-ray	6	SR	[23]
lrd32	苗期开始出现棕色类病斑,到种子成熟开始逐渐消失	γ-ray	-	DR	[8]
lrd39	出现棕色病斑直至种子成熟	γ-ray	-	SR	[8]
lrd40	苗期开始出现棕色类病斑,到种子成熟开始逐渐消失	γ-ray	-	SR	[8]
lrd42	叶片出现棕色稀疏类病斑	γ-ray	-	SR	[8]
Blm	叶片上出现褐色或者白色的类病斑	MNU	-	SR	[24]
lmi	最先出现在叶片尖部,最后蔓延整个叶片	γ-ray	8	SR	[25]
lmm1	4叶期最初出现褐色类病斑,接着类病斑扩散蔓延至整个叶片	EMS	-	SR	[26]
ttm1	幼叶开花早期,叶片上出现黄褐色病斑	Tos17	5	SR	[27]
OsLSD1	叶片上产生褐色类病斑	cDNA	8	SR	[28]
NPR1	叶片出现红棕色类病斑	Overexpression	-	SD	[29]
ncr1	叶片出现褐色类病斑,叶片发黄	FN	-	SR	[30]
lbsl1	播种后10~14d,叶片上出现淡褐色类病斑,然后扩散蔓延至整个叶片	EMS	6	SR	[31]
突变体 Mutant	病斑表型特征 Phenotype of lesion	突变体来源 Origin	染色体 Chr.	遗传方式^a	文献 Ref.
HM47	在幼苗叶片上会出现比较密集的棕褐色或者黑色类病斑	EMS	4	SR	[32]
splNF4050-8	叶片上首先出现黄色类病斑,生长后期类病斑变成棕褐色	Tissue	7	SR	[33]
c5	播种3~4周叶尖出现褐色小斑点,后蔓延到叶片和叶脉,呈深褐色	EMS	5	SR	[34]
LM3	叶片上出现褐色类病斑	γ-ray	3	SR	[35]
lms1	苗期首先出现棕黄色类病斑,随着叶片生长类病斑变成黄白色条斑	γ-ray	6	SR	[36]
OsACDR1	叶片出现棕色病斑,过量表达导致植株早衰	cDNA	3	SR	[37]
RLIN1	首先在幼叶和叶鞘上出现条纹状或者点状病斑,然后扩散蔓延	T-DNA	4	SR	[38]
NLS1	叶片尖部出现黄色类病斑	T-DNA	11	SR	[39]
OsLMS	扩散型类病斑	EMS	2	SR	[40]
C23	3叶期叶片首现淡黄色病斑,后病斑明显增多最终将蔓延到整个植株	EMS	12	SR	[41]
g303/g342	播种4~5周叶片现橘黄色类病斑,后扩散成中心枯黄边缘黄褐色的不规则斑块	γ-ray	12	SR	[42]
lmes1	叶片尖部首先出现黄色类病斑,然后蔓延到整个叶片,呈枯黄色	γ-ray	7	SR	[43]
splHM143	播种后3周,叶片出现褐色类病斑并向四周扩散,类病斑伴随整个生育期	EMS	4	SR	[44]
lmm4	分蘖期后期植株上部叶片从叶尖开始出现红褐色类病斑,随后扩散至整个叶片	EMS	5	SR	[45]
hm197	播种65d后,叶片尖部开始出现褐色病斑并扩散蔓延整个叶片	DEB	4	SR	[46]
wy3	苗期开始叶片出现白色类病斑,分蘖期较为明显,然后扩散到整个叶片	Spontaneous	2	SR	[47]
hm113	播种后3周叶片上出现褐色类病斑,始穗期类病斑扩散到叶鞘	EMS	7	SD	[48]
hm83	叶片上出现大小不一的红褐色类病斑	EMS	1	SR	[49]

突变体 Mutant	病斑表型特征 Phenotype of lesion	突变体来源 Origin	染色体 Chr.	遗传方式^a	文献 Ref.
spl1	播种后15~20d,叶片或者叶鞘出现橙黄色类病斑并扩散到整个叶片	Spontaneous	12	SR	[6]
spl2	苗期叶片和茎表面出现不明显的变色,分蘖期变色更加明显	Spontaneous	2	SR	[11]
spl3	在分蘖期到抽穗期,叶片上出现较小的红色类病斑	γ-ray	3	SR	[11]
spl4	叶片上零散出现红棕色类病斑	γ-ray	6	SR	[11]
spl5	在分蘖期到抽穗期,叶片上出现较小的红色类病斑	γ-ray	7	SR	[12]
spl6	分蘖期叶片上出现较大的棕色类病斑	MNU	1	SR	[13]
spl7	在分蘖期到抽穗期,叶片上出现较小的红色类病斑	γ-ray	5	SR	[14]
spl8	分蘖期后,整个叶片表面都会出现红棕色类病斑	MNU	5	SR	[11]
spl9	植株花期后,在叶片和茎上会有较小的黑棕色类病斑	γ-ray	7	SR	[11]
spl10	在叶片上有黄色或棕色的类病斑	MNU	10	SR	[11]
spl11	分蘖期后出现铁锈样的类病斑,然后到成熟期后,病斑蔓延到整个叶片	EMS	12	SR	[15]
spl12	苗期,叶片枯萎且表现出黄绿色	MNU	-	SD	[16]
spl13	播种后10周后,植株叶片出现几毫米长的棕色类病斑	MNU	-	SD	[16]
spl14	播种后10周,叶片上出现暗棕色类病斑	MNU	-	SR	[16]
spl15	播种后3~4周,出现5~10mm的淡棕色类病斑	MNU	-	SD	[16]
spl16	叶片上出现小的黑色类病斑	DEB	-	SR	[10]
spl17	叶片上出现小的棕褐色类病斑	DEB	-	SR	[10]
spl18	叶片上出现小的棕褐色类病斑	T-DNA	10	SD	[10]
spl19	叶片上出现棕黄色类病斑,叶片变卷曲,变窄	DEB	-	SR	[10]
spl20	叶片上出现白色类病斑	FN	-	SR	[10]
spl21	叶片上出现棕黄色类病斑	DEB	12	SR	[10]
spl22	叶片上出现黑色类病斑	DEB	-	SR	[10]
spl23	叶片上出现黄色类病斑	DEB	-	SR	[10]
spl24	叶片上出现多而密集的黑色斑点	DEB	-	SD	[10]
spl25	叶片上出现深绿色斑点	DEB	-	SR	[10]
spl26	基因纯合致死,叶片出现红褐色类病斑	DEB	-	SD	[10]
spl27	基因纯合致死,叶片出现棕黄色类病斑	DEB	-	SD	[10]
spl28	植株产生类病斑的同时伴随植株的严重早衰,病斑呈黄褐色	MNU	1	SR	[17]
spl30	叶片出现红褐色类病斑并逐渐扩散	EMS	3	SR	[18]
spl30(t)	叶片出现红棕色类病斑	Transgene	12	SR	[18]
spl31	叶片出现棕色类病斑	EMS	12	SR	[19]
spl32	叶片边缘出现棕黄色类病斑	-	11	SD	[20]
bl1	叶片上首先出现棕色病变斑点,然后蔓延到穗上	Spontaneous	1	SR	[11]
bl2	叶片出现黑色类病斑并逐渐蔓延扩散	Spontaneous	2	SR	[11]
bl3	叶片上出现黑色类病斑,植株成熟后类病斑会蔓延到颖壳	Spontaneous	6	SR	[11]
bl4	植株上大范围出现大的棕色类病斑	32P	3	SR	[11]
bl5	叶片上出现很小的点状棕色类病斑	32P/X-ray	-	SR	[11]
bl6	叶片上出现较大的棕色类病斑	32P	-	SR	[11]
cdr1	叶片上出现类病斑,为起始型	MNU	-	SR	[21]
cdr2	起始型类病斑	MNU	-	SR	[21]
cdr3	基因纯合致死	MNU	-	SD	[21]
Fgl	叶片伸长过程中,叶片出现黄色或者白色的类病斑	Spontaneous	10	SR	[22]
Zn	叶片上出现黄色类病斑	γ-ray	6	SR	[23]
lrd32	苗期开始出现棕色类病斑,到种子成熟开始逐渐消失	γ-ray	-	DR	[8]
lrd39	出现棕色病斑直至种子成熟	γ-ray	-	SR	[8]
lrd40	苗期开始出现棕色类病斑,到种子成熟开始逐渐消失	γ-ray	-	SR	[8]
lrd42	叶片出现棕色稀疏类病斑	γ-ray	-	SR	[8]
Blm	叶片上出现褐色或者白色的类病斑	MNU	-	SR	[24]
lmi	最先出现在叶片尖部,最后蔓延整个叶片	γ-ray	8	SR	[25]
lmm1	4叶期最初出现褐色类病斑,接着类病斑扩散蔓延至整个叶片	EMS	-	SR	[26]
ttm1	幼叶开花早期,叶片上出现黄褐色病斑	Tos17	5	SR	[27]
OsLSD1	叶片上产生褐色类病斑	cDNA	8	SR	[28]
NPR1	叶片出现红棕色类病斑	Overexpression	-	SD	[29]
ncr1	叶片出现褐色类病斑,叶片发黄	FN	-	SR	[30]
lbsl1	播种后10~14d,叶片上出现淡褐色类病斑,然后扩散蔓延至整个叶片	EMS	6	SR	[31]
突变体 Mutant	病斑表型特征 Phenotype of lesion	突变体来源 Origin	染色体 Chr.	遗传方式^a	文献 Ref.
HM47	在幼苗叶片上会出现比较密集的棕褐色或者黑色类病斑	EMS	4	SR	[32]
splNF4050-8	叶片上首先出现黄色类病斑,生长后期类病斑变成棕褐色	Tissue	7	SR	[33]
c5	播种3~4周叶尖出现褐色小斑点,后蔓延到叶片和叶脉,呈深褐色	EMS	5	SR	[34]
LM3	叶片上出现褐色类病斑	γ-ray	3	SR	[35]
lms1	苗期首先出现棕黄色类病斑,随着叶片生长类病斑变成黄白色条斑	γ-ray	6	SR	[36]
OsACDR1	叶片出现棕色病斑,过量表达导致植株早衰	cDNA	3	SR	[37]
RLIN1	首先在幼叶和叶鞘上出现条纹状或者点状病斑,然后扩散蔓延	T-DNA	4	SR	[38]
NLS1	叶片尖部出现黄色类病斑	T-DNA	11	SR	[39]
OsLMS	扩散型类病斑	EMS	2	SR	[40]
C23	3叶期叶片首现淡黄色病斑,后病斑明显增多最终将蔓延到整个植株	EMS	12	SR	[41]
g303/g342	播种4~5周叶片现橘黄色类病斑,后扩散成中心枯黄边缘黄褐色的不规则斑块	γ-ray	12	SR	[42]
lmes1	叶片尖部首先出现黄色类病斑,然后蔓延到整个叶片,呈枯黄色	γ-ray	7	SR	[43]
splHM143	播种后3周,叶片出现褐色类病斑并向四周扩散,类病斑伴随整个生育期	EMS	4	SR	[44]
lmm4	分蘖期后期植株上部叶片从叶尖开始出现红褐色类病斑,随后扩散至整个叶片	EMS	5	SR	[45]
hm197	播种65d后,叶片尖部开始出现褐色病斑并扩散蔓延整个叶片	DEB	4	SR	[46]
wy3	苗期开始叶片出现白色类病斑,分蘖期较为明显,然后扩散到整个叶片	Spontaneous	2	SR	[47]
hm113	播种后3周叶片上出现褐色类病斑,始穗期类病斑扩散到叶鞘	EMS	7	SD	[48]
hm83	叶片上出现大小不一的红褐色类病斑	EMS	1	SR	[49]

基因 Gene	功能及调节途径 Function and pathway
Spl5	编码特定的剪接因子3b亚基3(SF3b3),并参与调节细胞死亡和抗性反应。它是通过调节植物中的RNA剪接来负调节细胞死亡和抗性反应的
Spl7	存在相似于热应激转录因子(HSF)的ORF,通过转基因分析得知spl7编码的HSF蛋白在一定的环境下对植物细胞的死亡起负调控作用
Spl11	具有依赖于完整U-box结构域的E3泛素连接酶活性的特点,在控制植物细胞死亡和防御中起着负调控的作用
Spl18	该突变基因控制的蛋白质OsAT1与烟草中过敏反应诱导的酰基转移酶序列相似,OsAT1在突变体spl18中表达水平很高,过度表达导致类病变出现,同时也表现出对稻瘟病抗性的增强
Spl28	早期发育是正常的,但是在开花后spl28突变体中叶绿素含量、可溶性蛋白质含量以及光合作用效率显著降低,而且在病斑周围的有高浓度的活性氧(ROS)、胼胝质和酚类化合物。spl28编码高尔基体上的网格蛋白相关受体蛋白复合体1(AP1)亚单位µ1(AP1M1),表现出对稻瘟病和白叶枯病的抗性增强
OsPti1a	突变基因编码质膜蛋白激酶,质膜蛋白激酶的过度表达会抑制突变体的表型,对水稻的防御反应和抗病性起负调控作用
OsNPR1	在SA介导下OsNPR1可以让水稻获得抗性,OsNPR1的过度表达使突变体拥有对白叶枯病的抗性
OsACDR1	又名OsEDR1, 其转录会促进乙烯的合成,乙烯反过来又抑制了水杨酸(SA)和茉莉酸(JA)相关的防御信号
OsSSI2	突变基因OsSSI2控制脂肪酸脱氢酶活性,该酶的表达水平提高会导致水稻出现类病变症状,延缓生长,同时水稻的抗病性增强,因此OsSSI2在水稻防御反应的过程中也是起着负调控的作用
Rls1	又名lrd43,是隐性等位基因,参与叶片衰老的过程,其编码一个含有N端核苷酸结合位点(NB)结构域和 C 端ARM(armadillo)结构域的蛋白
Nls1	nls1突变基因伴随防御反应组成性表达,包括广泛的细胞死亡,过量的过氧化氢和水杨酸（SA）积累,导致对病原的抗性增强。另外nls1编码的蛋白具有CC-NB-LRR结构,参与植物的防御反应
Rlin1	Rlin1突变基因编码卟啉原III氧化酶,其病变的形成依赖于活性氧积累
OsLMS	编码羧基末端结构域(CTD)磷酸酶结构域和含两个双链RNA结合域(dsRBM）蛋白质产物,它控制植物应激反应和生长发育,是水稻胁迫反应的负调节因子
SL	它在细胞色素P450单加氧酶家族中编码CYP71P1蛋白,重组SL蛋白显示色胺5羟化酶活性,并催化色胺转化为5-羟色胺,5-羟色胺在水稻先天免疫中起重要作用
OsLSD1	OsLSD1是拟南芥LSD1在水稻中的功能同源物,含有保守的锌指结构域,OsLSD1的表达是光诱导或暗抑制的,对水稻防卫反应和细胞程序化死亡起负调控作用
OsHPL3	编码脂氢过氧化物裂解酶,OsHPL3活性调节了JA生物合成和信号传导的活化,激活JA控制的防御反应,从而影响水稻的抗病性
Spl29	SPL29编码UDP-N-乙酰葡糖胺焦磷酸化酶(UAP1),突变体spl29缺乏UAP活性。UAP1参与调控水稻叶片衰老和防御反应
LMR	LMR编码与各种AAA-类型蛋白相关的ATP酶,LMR是水稻细胞死亡的负调节剂,同时表现出对水稻稻瘟病和白叶枯病抗性增强
lmm5	通过增加防御相关基因的表达和过氧化氢的积累,表现出对水稻稻瘟病和白叶枯病的抗性增强
Spl33	Spl33编码由非功能性锌指结构域和三个功能性EF-Tu结构域组成的真核翻译延伸因子1α（eEF1A）样蛋白,实验表明活性氧的积累可能是spl33突变体中细胞死亡的原因,而多种先天免疫相关受体基因和信号分子的失控激活可能是spl33抗病性增强的原因。进一步分析得知eEF1A样蛋白参与细胞程序性死亡,并提供了与水稻防御反应相关的途径
OsDRP1E	OsDRP1E通过控制线粒体结构和细胞色素c的释放来负调节PCD

基因 Gene	功能及调节途径 Function and pathway
Spl5	编码特定的剪接因子3b亚基3(SF3b3),并参与调节细胞死亡和抗性反应。它是通过调节植物中的RNA剪接来负调节细胞死亡和抗性反应的
Spl7	存在相似于热应激转录因子(HSF)的ORF,通过转基因分析得知spl7编码的HSF蛋白在一定的环境下对植物细胞的死亡起负调控作用
Spl11	具有依赖于完整U-box结构域的E3泛素连接酶活性的特点,在控制植物细胞死亡和防御中起着负调控的作用
Spl18	该突变基因控制的蛋白质OsAT1与烟草中过敏反应诱导的酰基转移酶序列相似,OsAT1在突变体spl18中表达水平很高,过度表达导致类病变出现,同时也表现出对稻瘟病抗性的增强
Spl28	早期发育是正常的,但是在开花后spl28突变体中叶绿素含量、可溶性蛋白质含量以及光合作用效率显著降低,而且在病斑周围的有高浓度的活性氧(ROS)、胼胝质和酚类化合物。spl28编码高尔基体上的网格蛋白相关受体蛋白复合体1(AP1)亚单位µ1(AP1M1),表现出对稻瘟病和白叶枯病的抗性增强
OsPti1a	突变基因编码质膜蛋白激酶,质膜蛋白激酶的过度表达会抑制突变体的表型,对水稻的防御反应和抗病性起负调控作用
OsNPR1	在SA介导下OsNPR1可以让水稻获得抗性,OsNPR1的过度表达使突变体拥有对白叶枯病的抗性
OsACDR1	又名OsEDR1, 其转录会促进乙烯的合成,乙烯反过来又抑制了水杨酸(SA)和茉莉酸(JA)相关的防御信号
OsSSI2	突变基因OsSSI2控制脂肪酸脱氢酶活性,该酶的表达水平提高会导致水稻出现类病变症状,延缓生长,同时水稻的抗病性增强,因此OsSSI2在水稻防御反应的过程中也是起着负调控的作用
Rls1	又名lrd43,是隐性等位基因,参与叶片衰老的过程,其编码一个含有N端核苷酸结合位点(NB)结构域和 C 端ARM(armadillo)结构域的蛋白
Nls1	nls1突变基因伴随防御反应组成性表达,包括广泛的细胞死亡,过量的过氧化氢和水杨酸（SA）积累,导致对病原的抗性增强。另外nls1编码的蛋白具有CC-NB-LRR结构,参与植物的防御反应
Rlin1	Rlin1突变基因编码卟啉原III氧化酶,其病变的形成依赖于活性氧积累
OsLMS	编码羧基末端结构域(CTD)磷酸酶结构域和含两个双链RNA结合域(dsRBM）蛋白质产物,它控制植物应激反应和生长发育,是水稻胁迫反应的负调节因子
SL	它在细胞色素P450单加氧酶家族中编码CYP71P1蛋白,重组SL蛋白显示色胺5羟化酶活性,并催化色胺转化为5-羟色胺,5-羟色胺在水稻先天免疫中起重要作用
OsLSD1	OsLSD1是拟南芥LSD1在水稻中的功能同源物,含有保守的锌指结构域,OsLSD1的表达是光诱导或暗抑制的,对水稻防卫反应和细胞程序化死亡起负调控作用
OsHPL3	编码脂氢过氧化物裂解酶,OsHPL3活性调节了JA生物合成和信号传导的活化,激活JA控制的防御反应,从而影响水稻的抗病性
Spl29	SPL29编码UDP-N-乙酰葡糖胺焦磷酸化酶(UAP1),突变体spl29缺乏UAP活性。UAP1参与调控水稻叶片衰老和防御反应
LMR	LMR编码与各种AAA-类型蛋白相关的ATP酶,LMR是水稻细胞死亡的负调节剂,同时表现出对水稻稻瘟病和白叶枯病抗性增强
lmm5	通过增加防御相关基因的表达和过氧化氢的积累,表现出对水稻稻瘟病和白叶枯病的抗性增强
Spl33	Spl33编码由非功能性锌指结构域和三个功能性EF-Tu结构域组成的真核翻译延伸因子1α（eEF1A）样蛋白,实验表明活性氧的积累可能是spl33突变体中细胞死亡的原因,而多种先天免疫相关受体基因和信号分子的失控激活可能是spl33抗病性增强的原因。进一步分析得知eEF1A样蛋白参与细胞程序性死亡,并提供了与水稻防御反应相关的途径
OsDRP1E	OsDRP1E通过控制线粒体结构和细胞色素c的释放来负调节PCD