木栓质组成成分、组织化学特性及其生物合成研究进展

doi:10.11983/CBB16068

植物学报 ›› 2017, Vol. 52 ›› Issue (3): 358-374.DOI: 10.11983/CBB16068

木栓质组成成分、组织化学特性及其生物合成研究进展

韩雪源, 茅林春^*()

浙江大学食品科学与营养系, 农业部农产品采后重点实验室, 浙江省农产品加工重点实验室, 杭州 310058

收稿日期:2016-04-01 接受日期:2016-06-24 出版日期:2017-05-01 发布日期:2017-05-27
通讯作者: 茅林春
作者简介:
# 共同第一作者
基金资助:
国家自然科学基金(No.31372113)

Research Progress on Constituents, Histochemical Characteristics and Biosynthesis of Suberin

Xueyuan Han, Linchun Mao^*

Zhejiang Key Laboratory of Agro-Food Processing, Key Laboratory of Postharvest Handling Agro-Products for Ministry of Agriculture, Department of Food Science and Nutrition, Zhejiang University, Hangzhou 310058, China

Received:2016-04-01 Accepted:2016-06-24 Online:2017-05-01 Published:2017-05-27
Contact: Mao Linchun
About author:
# Co-first authors

摘要/Abstract

摘要： 木栓质是一种植物次生代谢产物, 是以甘油为基础的多聚脂类和多聚酚类物质的聚合物, 定位于细胞壁和质膜之间, 主要分布在根皮层和茎的次生边界组织中, 可以降低细胞水分和营养物质的外流, 限制病原体的入侵, 阻碍有毒气体向内扩散。近年来, 随着人们对果蔬贮藏和植物抗性的关注, 对木栓质的研究越来越深入, 尤其是代谢相关酶及基因功能研究。该文系统阐述木栓质组织化学、生物合成及其相关酶和基因的研究动态, 介绍木栓质的生理功能, 总结木栓质组成物质的转运、聚合及其堆积调控等, 以期为木栓质的深入研究和开发利用提供参考。

Abstract: Located between the cell wall and plasma membrane as a secondary metabolite, suberin typically distributes in rhizodermis and the boundary tissue of stems. Based on glycerol, suberin is a heteropolymer composed of polya- liphatics and polyaromatics, and it could slow the outflow of water and nutrient substance, limit pathogen invasion and prevent toxic gas from diffusing to plants. Recently, with people’s focus on the storage and processing of fruits and vegetables, as well plant resistance, the research in suberin is increasing, especially in the aspects of metabolic enzymes and corresponding genes and the metabolite’s function. In this paper, we elaborate the research progress in suberin histochemistry, the biosynthesis pathway as well related enzymes and genes. We introduce recent advances in the transport of suberin components intracellularly and to the cell wall, polymer assembly, and the regulation of suberin deposition and present the research development of suberin physiological function. This research is expected to provide significant information for further research and application of suberin.

韩雪源, 茅林春. 木栓质组成成分、组织化学特性及其生物合成研究进展. 植物学报, 2017, 52(3): 358-374.

Xueyuan Han, Linchun Mao. Research Progress on Constituents, Histochemical Characteristics and Biosynthesis of Suberin. Chinese Bulletin of Botany, 2017, 52(3): 358-374.

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https://www.chinbullbotany.com/CN/Y2017/V52/I3/358

图/表 2

图1 木栓质在根不同发育阶段的堆积位点(改自Vishwanath et al., 2015)图中所示为典型双子叶植物(例如拟南芥)根的横切面。(A) 内胚层发育过程中细胞壁的初生生长阶段, 木栓质片层在其内表面有所积累(黄色), 凯氏带(红色)在内皮层细胞间的结合点处积累(ML: 中间层; PW: 初生细胞壁; SL: 木栓质片层; PM: 细胞质膜; Cy: 细胞质); (B) 次生生长阶段, 木栓质堆积于木栓组织/栓化细胞(周皮)中

Figure 1 Sites of suberin deposition in different developmental stages of roots (modified from Vishwanath et al., 2015)The diagrams show the root cross-sections of typical dicot plants (e.g. Arabidopsis thaliana). (A) Primary growth stage of cell wall during endodermis development with suberin lamellae deposition in the inner face of endodermal cell walls (yellow) and deposition of Casparian bands (red) localized at endodermal cell-cell junctions (ML: Middle lamellae; PW: Primary cell wall; SL: Suberin lamellae; PM: Plasma membrane; Cy: Cytoplasm); (B) Secondary growth stage with suberin deposition in the phellem/cork cells (periderms)

图2 木栓质合成及其运输至细胞壁的路径概述脂肪酸合成发生在细胞质内, 之后被脂肪酸-CoA连接酶(FACL)和长链酰基-CoA合成酶(LACSs)活化, 生成脂肪酰CoA类物质, 然后被一系列木栓质合成酶修饰。脂肪酸延长酶(FAE)复合体调控脂肪酰延长, 产生超长链脂肪酸(VLCFAs); 脂肪酰还原酶(FARs)还原酰基, 生成脂肪醇类和α, ω-二醇; 细胞色素P450酶类(CYPs)氧化脂肪酰, 生成ω-羟基脂肪酸(ω-OHs)和α, ω-二羧酸(DCAs); ω-OHs可被细胞色素P450s进一步氧化成DCAs; ω-OHs和DCAs被甘油-3-磷酸酰基转移酶(GPATs)酯化成sn-2单酰基甘油酯。ATP-结合盒(ABC)转运蛋白参与木栓质单体穿过质膜的运输。多酯合酶(PS)可能用其它木栓质单体延伸sn-2单酰基甘油, 使所有成分聚集, 产生高分子质量聚酯。在苯丙氨酸解氨酶(PAL)等酶作用下, 苯丙烷代谢途径生成香豆酸、咖啡酸和阿魏酸, 它们在酰基连接酶等作用下, 再生成阿魏酰-CoA。阿魏酰-CoA在脂肪木栓质阿魏酰转移酶(ASFT)作用下, 与脂肪醇、α, ω-二醇和单酰基甘油酯连接, 再生成相应的酯类物质。

Figure 2 Overview of the suberin biosynthetic pathway with subsequent transport to the cell wallFatty acids are synthesed in the cytoplasm, activated into fatty acyl-CoAs by fatty acid-CoA ligase (FACL) and long chain acyl-CoA synthetases (LACSs), and then modified by series suberin biosynthetic enzymes. Fatty acyl elongation is controlled via the fatty acid elongation (FAE) complex producing very long chain fatty acids (VLCFAs); acyl reduction by fatty acyl reductases (FARs) producing fatty alcohol and α, ω-diols; fatty acyl oxidation by cytochrome P450 enzymes (CYPs) producing ω-hydroxy fatty acids (ω-OHs) and α, ω-dicarboxylic acids (DCAs); ω-OHs could be also further oxidized to DCAs by P450s; and esterification of ω-OHs and DCAs by glycerol 3-phosphate acyltransferases (GPATs) producing sn-2 monoacylglycerols. ATP-binding- cassette (ABC) transporters are involved in transport of suberin monomers across the plasma membrane. Polyester synthase(s) (PS) may extend sn-2 monoacylglycerols with other suberin monomers to eventually gather into high molecular weight polyesters. With the catalytic action of phenylalnine ammonialyase (PAL) and other related enzymes, phenylalanine metabolic pathways provide coumaric, caffeic, and ferulic acids, which then are transformed to feruloyl-CoA by acyltransferase. Moreover, feruloyl-CoA is linked with fatty alcohols, α, ω-diols and monoacylglycerols through aliphatic suberin feruloyl transferase (ASFT), to finally produce esters.

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木栓质组成成分、组织化学特性及其生物合成研究进展

Research Progress on Constituents, Histochemical Characteristics and Biosynthesis of Suberin

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