糖信号及其在植物响应逆境胁迫中的作用

doi:10.13560/j.cnki.biotech.bull.1985.2021-1289

摘要/Abstract

摘要：

糖不仅是植物细胞的碳源、能源和结构物质,也是一种信号分子,在植物生长发育及响应逆境胁迫中起重要作用。非生物逆境如高温、低温、干旱、盐渍和重金属胁迫是限制作物产量的主要胁迫因子,糖作为信号分子在植物响应这些胁迫因子中的确切机理,尚未清楚。基于植物中糖信号转导途径及其在植物耐逆性形成中作用的最新研究进展,归纳了植物中分别依赖己糖激酶（HXK）、G蛋白信号1调节子（RGS1）、糖酵解（EMP）和磷酸戊糖途径（PPP）的糖信号转导途径,讨论了糖信号在植物耐逆性包括耐热性、耐冷性、耐旱性、耐盐性和重金属胁迫耐性形成中的作用,最后展望了糖信号在植物生物学领域的研究方向。

关键词: 糖信号, 耐热性, 耐冷性, 耐旱性, 耐盐性, 重金属胁迫耐性

Abstract:

Sugar is not only carbon and energy sources as well as structural material in plant cells,but also a signaling molecule,which plays a key role in plant growth,development,and response to environmental stresses. Abiotic stresses,such as high temperature,low temperature,drought,salinity,and heavy metal,are the major stress factors limiting crop yield. The exact mechanism of sugar,as signaling molecule,in plants response to these stresses still remains unclear. Based on the current advance in sugar signaling and its role in the formation of stress tolerance in plants,the hexose kinase（HXK）-,regulator of G protein signaling 1（RGS1）-,glycolysis（EMP）-,and pentose phosphate pathway（PPP）-dependent signaling pathways were separately summarized;and the role of sugar signaling in the developing of plant stress tolerance,including heat,cold,drought,salt,and heavy metal stress tolerance was discussed. In addition,the future research direction of H₂S signaling in the field of plant biology was prospected.

Key words: sugar signaling, heat tolerance, cold tolerance, drought tolerance, salt tolerance, heavy metal stress tolerance

陈宏艳, 李小二, 李忠光. 糖信号及其在植物响应逆境胁迫中的作用[J]. 生物技术通报, 2022, 38(7): 80-89.

CHEN Hong-yan, LI Xiao-er, LI Zhong-guang. Sugar Signaling and Its Role in Plant Response to Environmental Stress[J]. Biotechnology Bulletin, 2022, 38(7): 80-89.

图/表 3

图1 植物糖信号转导途径植物可分别通过依赖己糖激酶（HXK）、G蛋白信号1调节子（RGS1）、糖酵解（EMP）和磷酸戊糖途径（PPP）的信号途径转导糖信号,继而上调叶绿素a/b结合蛋白1（CAB1）、质体蓝素（PC）和核酮糖1, 5-二磷酸羧化/加氧酶（Rubisco）、查尔酮合酶（CHS）、苯丙氨酸裂解酶（PAL）、腺苷二磷酸-葡萄糖焦磷酸化酶（AGPase）、天冬酰氨合成酶1（AS1）、蛋白S6激酶（S6K）、自噬相关蛋白（ATG）、NO3-转运蛋白（NTR）、SO42-转运蛋白（SULTR）等的基因表达,最终整合生长发育及逆境响应。依赖HXK、RGS1、EMP和PPP的信号途径可分别被2-脱氧甘露糖（Man）、2-脱氧葡萄糖（2-DG）、3-O-甲基葡萄糖（3-OMG）和6-脱氧葡萄糖（6-DG）所促进,被甘露庚酮糖（MHP）、葡萄糖胺（GCN）、苏拉明（SUR）、AZD-8055、葡萄糖-6-磷酸（G6P）和6-氨基烟酰胺（6-AN）所抑制。（+）表示促进,（├）表示抑制

Fig. 1 Sugar signaling pathways in plants Plants can transduce sugar signaling by hexokinase（HXK）-,regulator of G protein signaling1（RGS1）-,glycolysis（EMP）-,and pentose phosphate pathway（PPP）-dependent signaling pathways,which in turn up-regulate gene expression of chlorophyll a/b-binding protein1（CAB1）、plastocyanin（PC）,ribulose-1, 5-bisphosphate carboxylase/oxygenase（Rubisco）,chalcone synthase（CHS）,phenylalanine amonia-lyase（PAL）,adenosine diphosphate-glucose pyrophosphorylase（AGPase）,asparagine synthetase1（AS1）,protein S6 kinase（S6K）,autophagy-related protein（ATG）,nitrate transporter（NTR）,and sulfate transporter（SULTR）,and then integrate plant growth,development,and response to environmental stress. The HXK-,RGS1-,EMP- and PPP-dependent sugar signaling pathways can be activated by 2-deoxymannose（Man）,2-deoxyglucose（2-DG）,3-O-methyl-D-glucose（3-OMG）and 6-deoxyglucose（6-DG）,respectively,while inactivated by mannoheptulose（MHP）,glucosamine（GCN）,suramin（SUR）,AZD-8055,glucose-6-phosphate（G6P）and 6-aminonicotinamide（6-AN）,respectively.（+）represents promotion,while（├）represents inhibiton

图2 糖信号在植物响应耐逆胁迫中的作用高温、低温、干旱、盐渍、重金属胁迫及外源提供可溶性糖,均可增加植物体内可溶性糖内源水平,继而通过引发糖信号途径和/或直接作用,增强植物激素、热激蛋白、渗透调节系统、抗氧化系统、甲基乙二醛脱毒系统等活力,植物以此响应逆境胁迫

Fig. 2 Role of sugar signaling in the response of plants to environmental stress Heat,cold,drought,salt,and heavy metal stress as well as exogenous application of sugar increase the level of endogenous sugar,and which triggers sugar signaling pathways and/or exerts direct action,followed by enhancement in the activity of plant hormone,heat shock proteins,osmoregulation system,antioxidant system,and methylglyoxal（MG）detoxification system,thus plants respond to environmental stress

图3 植物细胞中糖的运输植物细胞可通过定位于质膜上的蔗糖运输蛋白（SUT）、糖运输蛋白（STP）、糖输出蛋白（SWEET）、蔗糖输入蛋白（U1）和己糖输入蛋白（U2）,以及定位于液泡膜上的蔗糖运输蛋白（SUT）、液泡葡萄糖运输蛋白（VGT）、SWEET、蔗糖输入蛋白（V1）和己糖输入蛋白（V2）、早期响应脱水蛋白类似蛋白（ERDL）、液泡膜蔗糖运输蛋白（TST）、液泡膜单糖运输蛋白（TMT）等转运蛋白运输糖,以响应生长发育及逆境胁迫

Fig.3 Sugar transport in plant cells Plant cells can transport sugar by sucrose transporter（SUT）,sugar transport protein（STP）,sugar will eventually be exported transporter（SWEET）,sucrose import protein（U1）,and hexose import protein（U2）locating in plasma membrane,as well as sucrose transporter（SUT）,vacuole glucose transporter（VGT）,SWEET,sucrose import protein（V1）,hexose import protein（V2）,early response dehydration protein-like（ERDL）,tonoplast sucrose transporter（TST）,and tonoplast monosaccharide transporter（TMT）locating in tonoplast,to response to growth,development environmental stress

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