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脯氨酸的亲水力极强,是构成蛋白质的唯一亚氨基酸。脯氨酸在植物中的作用和机制已得到广泛研究,除作为渗透调节物质外,脯氨酸还在清除细胞活性氧或作为信号分子调控植物细胞生长发育、增殖或死亡中发挥重要作用。现有研究结果表明,适当浓度的脯氨酸在细菌细胞中发挥重要功能。本文对细菌脯氨酸的合成、降解、在细胞内外的转运及功能进行综述。
Proline with strong hydrophilic property is the only sub-amino acid being used for protein synthesis. The function and mechanism for proline has been extensively studied in plant. Proline is not only an osmolyte, but it also has important roles in scavenging reactive oxygen species, acting as a signal molecule to regulate growth, development, proliferation and cell death in plant. Recent studies have demonstrated that a proper amount of proline plays important roles in bacterial cells. The biosynthesis, degradation, intracellular and extracellular transport as well as the function of proline produced by bacteria were reviewed in this paper.
L-脯氨酸,简称脯氨酸(proline,Pro),化学名称为吡咯烷酮羧酸,是构成蛋白质的20种基本氨基酸中唯一的亚氨基酸,亲水力极强,是一种天然的渗透保护剂和抗氧化剂[
在细菌中,澳大利亚学者Christian 1955年在
目前,已报道的脯氨酸合成途径主要有两条,根据起始物的不同分为谷氨酸(Glu)途径和鸟氨酸(Orn)途径[
以谷氨酸为起始物的脯氨酸合成途径首先在
细菌脯氨酸的合成与降解(谷氨酸途径)
Biosynthesis and degradation of proline in bacteria (glutamate pathway).
目前,已报道的比较确定的由鸟氨酸合成脯氨酸的途径有两条(
细菌脯氨酸的合成与降解(鸟氨酸途径)
Biosynthesis and degradation of proline in bacteria (ornithine pathway).
目前已知,δ-OAT是一种几乎存在于所有真核生物中的酶,可催化鸟氨酸向谷氨酸或脯氨酸的转化。在原核生物中,OAT (
由鸟氨酸合成脯氨酸的另一条途径是L-鸟氨酸被鸟氨酸环化脱氨酶(OCD)催化直接生成L-脯氨酸。该酶于1971年在梭状芽孢杆菌中发现[
脯氨酸代谢的中间产物P5C/GSA可被分解代谢和合成代谢途径共享,但脯氨酸的降解过程并非完全为其合成过程的逆反应,催化反应的酶亦不同。如
参与脯氨酸氧化的PRODH和P5CDH在真核生物和原核生物中均高度保守,所不同的是这两个酶是否融合为一个被称为脯氨酸利用A (PutA)的双功能酶。根据Tanner 2008年报道,脯氨酸降解酶可分为三类:第一类由单功能酶组成,即PRODH域和P5CDH域是分开的;第二类和第三类在同一条PutA多肽中均同时含有PRODH域和P5CDH域,二者的区别在于是否在N末端还含有一个RHH (ribbon-helix-helix) DNA结合域[
1982年,Meile等提出在革兰氏阴性铜绿假单胞菌PAO中,双功能脯氨酸降解酶的产生受脯氨酸诱导而不受P5C诱导,受柠檬酸盐和氮控制的降解物阻遏[
在革兰氏阳性枯草芽孢杆菌中,
现有研究结果表明,在大肠杆菌和沙门氏菌中的脯氨酸转运系统包括PutP、ProP和ProU。当菌株以脯氨酸为碳源或氮源时,PutP是其运输所必需,此时PutP可被外源脯氨酸诱导;ProP和ProU则用于运输渗透保护剂[
许多革兰氏阳性细菌脯氨酸的合成或降解则受渗透胁迫调控,在缺乏外源脯氨酸时,可通过增加脯氨酸的内源性合成应对渗透胁迫。与上述革兰氏阴性细菌一样,当革兰氏阳性枯草芽孢杆菌以脯氨酸作为碳或氮源时,其
Zaprasis等(2014年)发现PutP和OpuE系统同时失活时,枯草芽孢杆菌细胞仍然将脯氨酸作为营养物利用,推测在枯草芽孢杆菌中可能存在第三类脯氨酸转运系统。于是利用
革兰氏阴性大肠杆菌、鼠伤寒沙门氏菌、克雷伯菌及革兰氏阳性枯草芽孢杆菌均可以脯氨酸作为唯一氮源或碳源[
脯氨酸是一种有效的相容性溶质,可抵御高渗透压对细胞生理的不利影响[
在大肠杆菌中,脯氨酸作为一种热保护剂,在热应激过程中可减少蛋白质聚集[
现有研究结果表明,脯氨酸对胞内寄生菌的毒力发挥有重要作用。Smith等2001年的研究发现,脯氨酸生物合成途径中最后一个基因
除对胞内寄生菌外,脯氨酸对非胞内寄生病原菌的毒力发挥也有重要作用。现有研究结果证实,许多病原菌依赖脯氨酸作为关键呼吸底物。2003年,Nagata等研究发现,脯氨酸是幽门螺杆菌在人胃定殖期间优先选择的呼吸底物,在感染幽门螺杆菌患者的胃液中脯氨酸含量比未感染者高10倍[
脯氨酸可影响生物体的许多代谢过程,脯氨酸代谢在不同生物过程中具有不同的作用机制,包括细胞信号传导、应激保护和能量产生等[
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