植物胼胝质合成酶研究进展

doi:10.16420/j.issn.0513-353x.2020-0379

摘要/Abstract

摘要：

胼胝质是一种β-1,3-葡萄糖聚合物,植物在生长发育和受到生物、非生物胁迫过程中均会有胼胝质沉积。胼胝质合成酶又称β-1,3-葡聚糖合成酶类似物,通常以多亚基复合物形式控制胼胝质的合成。本文中对胼胝质合成酶的分离鉴定过程进行了梳理,对胼胝质合成酶复合物的亚基及其作用进行了描述,重点总结了转录因子、植物生长调节剂等对胼胝质合成酶基因表达的调控作用,并对胼胝质合成酶在花粉发育和韧皮部运输过程中的作用,对机械损伤、磷酸盐、金属离子等非生物胁迫,以及虫害、细菌和真菌等生物胁迫的应答反应进行了归纳;最后对胼胝质合成酶的研究方向提出展望,以期为解析胼胝质合成酶的调控机制提供参考,也为植物（特别是园艺植物）抗性基因的挖掘提供借鉴。

关键词: 胼胝质, 胼胝质合成酶, 生长发育, 胁迫响应, 调控机制

Abstract:

Callose is a β-1,3-glucan polymer,which is deposited during plant growth and development,biotic and abiotic stresses. Callose synthase,also known as β-1,3-glucan synthase like,usually controls the synthesis of callose in the form of multi-subunit complexes. The isolation and identification process of callose synthase,the subunits of the callose synthase complex and their roles are described in the review. The processes of callose synthase gene regulated by transcription factors,plant growth regulator and others are highlighted. In addition,the function of callose synthase under abiotic stresses（wound,phosphate,metal ions,etc.）and biotic stresses（pests,bacteria,fungi,etc.） are summarized. Finally,the research direction of callose synthase in future is discussed. The related content would be helpful for illuminating regulating mechanism of callose synthase and searching for resistance genes in plant（especially horticultural plants）.

Key words: callose, callose synthase, growth and development, stress response, regulation mechanism

中图分类号:

张庆雯, 王兆昊, 祁静静, 谢宇, 雷天刚, 何永睿, 陈善春, 姚利晓. 植物胼胝质合成酶研究进展[J]. 园艺学报, 2021, 48(4): 661-675.

ZHANG Qingwen, WANG Zhaohao, QI Jingjing, XIE Yu, LEI Tiangang, He Yongrui, CHEN Shanchun, YAO Lixiao. The Advances of Callose Synthase in Plant[J]. Acta Horticulturae Sinica, 2021, 48(4): 661-675.

图/表 3

表1 植物胼胝质合成酶基因和蛋白特征

Table 1 Gene and protein characteristics of plant callose synthase

物种 Species	数量 Number	开放阅读框/bp Open reading frame	内含子数 Intron number	外显子数 Exon number	编码氨基酸数 Amino acid number	相对分子质量/kD Relative molecular weight	文献 Reference
拟南芥 A rabidopsis thaliana	12	5 307 ~ 5 853	1 ~ 49	2 ~ 50	1 768 ~ 1950	205.00 ~ 226.00	Hong et al.2001a
甜橙Citrus sinensis	9/12	1 509 ~ 7 602	0 ~ 109	1 ~ 110	502 ~ 2533	186.53 ~ 581.38	Granato et al.2019;彭蕴等,2019
油菜 Brassica napus	32	1 812 ~ 6 915	0 ~ 54	1 ~ 55	603 ~ 2304	69.10 ~ 266.81	Liu et al.2018
葡萄Vitis vinifera	8	3 867 ~ 6 159	9 ~ 48	10 ~ 49	1 288 ~ 2052	149.96 ~ 237.77	Yu et al.2016
小立碗藓 Physcomitrella patens	12	5 268 ~ 5 901			1 755 ~ 1966		Schuette et al.2009
大麦 Hordeum vulgare	7	627 ~ 5 748			208 ~ 1915	23.50 ~ 218.00	Li et al.2003;Schober et al.2009
辣椒Capsicum annuum	9	5 265 ~ 6 393	2 ~ 51	3 ~ 52	1 754 ~ 2130		Sun et al.2018

图1 胼胝质合成酶复合物模型（Verma & Hong, 2002） SuSy:蔗糖合成酶;UGT:UDP-葡萄糖转移酶;Rop1:Rho样蛋白;ANN:膜联蛋白。

Fig. 1 Hypothetical model of the callose synthase complex（Verma & Hong, 2002） SuSy:Sucrose synthase;UGT:UDP-glucose transferase;Rop1:Rho-like protein;ANN:Annexin.

图2 调控胼胝质合成酶（CalS）的信号通路黄色为转录因子,绿色为植物生长调节剂,粉色为胼胝质合成酶。ARF7:生长素响应因子7;ARF17:生长素响应因子17;AFBs:生长素信号F-box蛋白;BPH14:褐飞虱抗性基因14;BAK1:BRI1-相关受体激酶1;BIK1:葡萄孢菌诱导的激酶1;CDPK:钙依赖蛋白激酶;CML41:钙调素类似蛋白41;COI1:冠状肽不敏感蛋白1;EDS1:易感性增强蛋白1;FLS2:鞭毛蛋白检测蛋白2;ICS1:分支酸合成酶1;I3CA:吲哚-3-羧酸;NCED3:9-顺式环氧类胡萝卜素双加氧酶3;NPR1:发病相关基因不表达子1;OCP3:阳离子型过氧化物酶3;PAD4:植物抗毒素不足蛋白4;PKL:染色质重塑因子;PR2:病程相关蛋白2;RBOH:呼吸爆发氧化酶同源物;SVL:短营养期类似蛋白;TTP:CCCH锌指蛋白（Tristetraproline）;TIR1:运输抑制剂响应蛋白1。

Fig. 2 Signal pathways to regulate callosesynthase（CalS） Yellow is the transcription factor,green is the plant growth regulator,pink is callosesynthase. ARF7:Auxin response factor 7;ARF17:Auxin response factor 17;AFBs:Auxin signaling F-Box protein;BPH14:Brown planthopper resistance 14;BAK1:Lrr-rk brassinosteroid receptor 1-associated kinase 1;BIK1:Botrytis-induced kinase 1;CDPK:Calcium-dependent protein kinases;CML41:Calmodulin-like 41;COI1:Coronatine-insensitive protein 1;EDS1:Enhanced disease susceptibility 1;FLS2:Flagellin sensing 2;ICS1:Isochorismate synthase 1;I3CA:Indole-3-caboxylic acid;NCED3:9′-cis-epoxycarotenoid dioxygenase;NPR1:Nonexpressor of pathogenesis-related genes 1;OCP3:Cationic peroxidase 3;PAD4:Phytoalexin deficient 4;PKL:Pickle;PR2:Pathogenesis-related protein 2;RBOH:Respiratory burst oxidase homologue;SVL:Short vegetative phase-like;TTP:Tristetraproline;TIR1:Transport inhibitor response 1.

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