植物油体研究进展

doi:10.11983/CBB16204

摘要/Abstract

摘要： 油体是生物体细胞中一种重要的细胞器结构, 由单层磷脂酸膜包裹中性脂肪酸形成, 膜上镶嵌有决定油体性质的多种膜蛋白。油体在能量储存、细胞生殖分化、抗病抗寒和发育调控等多种生命活动中起重要作用。该文对植物油体的结构、生物学功能、不同组织中油体的形成情况以及油体膜蛋白的研究进展进行了多方位概述和总结, 以期为后续研究提供有益的参考。

关键词: 油体, 油体结构, 生物学功能, 油体蛋白

Abstract: The oil body, covered with single layer phospholipid membrane, is an important organelle in eukaryotic organisms. Different oil-body proteins embedded in the membrane are crucial for the properties and functions of the oil body. The oil body plays important roles in energy reserve, cell proliferation and differentiation, disease and cold resistance and developmental regulation. In this review, we summarize recent research progress in the structure, biological function, and formation of the oil body in different tissues as well as membrane proteins embedded in the oil body, which will provide a useful reference for future research of oil bodies.

Key words: oil body, oil body architecture, biological function, oil body protein

胡佳, 刘春林. 植物油体研究进展. 植物学报, 2017, 52(5): 669-679.

Jia Hu, Chunlin Liu. Research Advances in Plant Oil Body. Chinese Bulletin of Botany, 2017, 52(5): 669-679.

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链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB16204

https://www.chinbullbotany.com/CN/Y2017/V52/I5/669

图/表 4

图1 油体形成模式图 (A) 油脂在内质网中通过脂肪酸合成酶DGAT (acyl-CoA: diacylglycerol acyltransferase)和ACAT (acyl-CoA:cholesterol acyltrans- ferase)合成后被细胞质面内质网膜包裹, 随着包裹体积的增大, 逐渐形成一个完全封闭的质体——油体, 随后油体与内质网分离或者继续黏结在一起, 通过内部三酰甘油的继续合成或者相互融合进行扩张(Walther and Farese, 2012)。(B) 合成的中性油脂进入内质网腔以后形成一种类似镜头的结构, 该结构为油体中间体。中间体的特殊结构被MHC家族的1类分子体(发卡状所示)识别后开始对中间体部位进行切割, 分别形成油体+MHC分子(右端所示)和磷脂酸+MHC分子(左端所示) 2种不同结构(Ploegh, 2007)。(C) 油体形成的蛋杯模式。内质网上存在的亲脂素能够将内质网膜上的中性油脂集合在一起, 形成一个油脂的核心, 然后内质网以蛋杯裹蛋的形式将含有磷脂单层膜的油脂核心包裹起来, 形成一个蛋杯样的结构。亲脂素存在部位与内质网上油脂部位密切相关。亲脂素在油体膜以及与油体膜接触的细胞质面内质网膜上大量积累, 质面内质网也有少量的亲脂素存在, 腔面内质网基本不含亲脂素(Robenek et al., 2006)。DAG: 二酰甘油; FA-CoA: 脂酰基辅酶A; TAG: 三酰甘油; SE: 甾醇酯; PC: 磷脂酰胆碱

Figure 1 Models of oil body formation (A) Neutral lipids are synthesized in the endoplasmic reticulum (ER) by the synthesis enzymes DGAT and ACAT and then covered with cytoplasmic face of the ER membrane. With the expansion of the membrane, a closed plastid called oil body is formed, and then the oil body discharges from the ER or remains adjacent with the ER membrane. The synthesis of neutral lipid in the oil body and the fusion of the oil body lead to the oil body expansion. (B) Neutral lipids enter the ER lumen and form a ‘lens’ structure between the two sheets of phospholipids which is called intermediate. Oil body intermediates are discharged by the recognition of bicelle (hairpin shown). The discharged parts either become part of oil body (right) or a structure composed solely of the class I MHC molecule and phospholipid (left). (C) The egg cup model of oil body synthesis. The adipophilin existing on the ER membrane can recruit the neutral lipid and form a lipid core, and then the ER membranes partially wrap the oil body and form an egg-cup like construct as egg cup wrapping egg. The location of adipophilin is closely related with the lipid existing. Large amounts of adipophilin accumulate on the membrane of oil body and the ER membrane contacting with oil body, low amounts at the cytoplasmic face of the ER, and almost does not exist at the lumen face of the ER (Robenek et al., 2006). DAG: Diacylglycerol; FA-CoA: Fatty acyl-coenzyme A; TAG: Triacylglycerol; SE: Sterol ester; PC: Phosphatidylcholine

图2 油体分子结构模式图(Walther and Farese, 2012)油体由磷脂酰胆碱、磷脂酰乙醇胺和磷脂酰肌醇等主要磷脂形成的单分子层膜包裹。磷脂膜上镶嵌着各类与油体合成和其生物学性质相关的油体蛋白。磷脂膜内为中性脂肪酸, 包括甾醇酯和三酰甘油等。

Figure 2 Schematic diagram of oil body architecture (Wal- ther and Farese, 2012)Oil body is covered with single layer membrane mainly con- stituted with phosphatidylcholine, phosphatidylethanolamine and phosphatidylinositol. The membrane is embedded with different kinds of oil body proteins that related with oil body synthesis and its bio-perspective. Inner the oil body, neural lipids including steroid esters and triacylglycerol are covered by the phospholipids membrane.

表1 主要油体膜蛋白及其特点一览表

Table 1 Mainly oil body membrane proteins and their properties

主要油体膜蛋白	主要存在部位	生物学功能	参考文献
油体蛋白(OLEs)	花粉和整个发育时期的种子	阻止/促进油体融合, 维持油体稳定/抗寒/保证种子正常萌发	Shimada et al., 2008; Krahmer et al., 2012
油体钙蛋白(Cals)	营养生长和生殖生长阶段	促进油体降解/维持油体稳定/抵抗生物胁迫/参与植物开花调节	Hara-Nishimura and Hatsugai, 2011; Shen et al., 2016
脂肪调控蛋白(SEIPIN)	营养生长和生殖生长阶段	控制油体体积和数量	Cai et al., 2015
油体协助蛋白1 (OBAP1)	种子胚	抑制油体降解?/保证种子正常萌发	Ghelis et al., 2008; López-Ribera et al., 2014
α-双加氧酶(α-DOX1)	营养生长和生殖生长阶段	抵抗真菌感染	Hara-Nishimura and Hatsugai, 2011
脂滴协助蛋白(LDAP)	营养生长和生殖生长阶段	参与植物昼夜循环/抗胁迫/TAG降解/种子萌发后生长调节	Gidda et al., 2016
糖依赖蛋白-1 (SDP1)	种子	种子萌发后降解TAG	Thazar-Poulot et al., 2015
NAD(P)H脱氢酶C1 (NDC1)	叶绿体	参与电子流途径和苯基醌的降解	Eugeni et al., 2011
固醇蛋白(Steroleosins)	种子和含油丰富的营养器官	参与甾醇酯的合成?	Shimada and Hara-Nishimura, 2010

图3 油体的形成及其生物学功能植物营养和生殖器官中形成的中性油脂在内质网上被单层磷脂酸膜包裹形成油体后, 在油体膜蛋白的辅助下参与植物多种生理活动。

Figure 3 The formation of oil body and its corresponding bio-functions The oil body, in both vegetative and reproductive organs, is formed by the neural lipid synthesized in the endoplasmic reticulum, and covered by the monolayer phospholipid and oil-body proteins. It play important roles in the physiological activities.

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