脯氨酸代谢与植物抗渗透胁迫的研究进展

脯氨酸被认为是植物和细菌内的一种相容渗透剂,有助于植物和细菌抵御渗透胁迫。本文就近年来有关植物体内脯氨酸合成和代谢、脯氨酸含量受渗透胁迫的影响情况、脯氨酸合成降解有关的酶及其基因、脯氨酸在细胞中的运输和定位、ABA与脯氨酸的诱导合成以及脯氨酸和植物抗渗透胁迫关系的研究进展作了简要综述。     

高等植物脯氨酸代谢研究进展

很多植物在胁迫条件下可以通过增加合成、减少降解而在体内累积大量脯氨酸,这对于调节渗透平衡、防止渗透胁迫对植物造成伤害、清除自由基、保护细胞结构具有重要意义。脯氨酸合成、降解相关酶的编码基因大都已经克隆到,但对脯氨酸在植物发育中的具体作用、胁迫条件下脯氨酸累积的分子机理了解还比较少。概述了植物控制脯氨酸合成、降解相关酶的编码基因的研究进展情况。     

脯氨酸在植物生长和非生物胁迫耐受中的作用

脯氨酸是生物界分布最广的渗透保护物质之一,干旱、高盐、高温及重金属等非生物胁迫条件都会导致植物体内脯氨酸含量的增加,其作用是防止渗透胁迫对植物造成的伤害、清除自由基,还可以作为氮、碳以及NADPH的重要来源。近年来,在转化脯氨酸代谢相关基因提高植物胁迫抗性方面也取得了很大进展。本文概要介绍了脯氨酸在植物生长和耐受非生物胁迫中的作用、与植物脯氨酸累积有关的信号转导、胁迫条件下脯氨酸的吸收和器官间的运输途径,以及通过转基因技术过量表达脯氨酸提高植物胁迫耐性的代谢工程的进展。     

细菌中脯氨酸的生物合成、降解及功能

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。     

渗透胁迫和外源脱落酸对梭梭幼苗生理特性的影响

水分、渗透胁迫和外源脱落酸（ＡＢＡ）对旱生植物梭梭幼苗的某些生理特性存在显著影响。梭梭幼苗有很强的吸水能力,渗透胁迫下脯氨酸含量增加,其细胞膜相对透性对渗透胁迫不敏感,在脱水过程中具有较强的持水能力,外源ＡＢＡ加强了梭梭幼苗的抗渗透胁迫和抗脱水的能力。     

植物体内脯氨酸累积和γ-谷氨酰磷酸合成活力的关系

渗透胁迫下植物体内脯氨酸合成加强,致使脯氨酸大量累积(汤章城 1984,Stewart和Hanson 1980,Dashek等1981)。在微生物中已经证实脯氨酸合成的关键酶为γ-谷氨酰激酶,并且已得到纯化酶(Baich 1969,Smith等1984),在高等植物中尚未完全证实这种酶的存在。Ericson等(1984)认为在高等植物中测定     

逆境条件下植物体内渗透调节物质的积累与活性氧代谢

本文介绍逆境胁迫下植物体内渗透调节物质的积累和作用,及其对活性氧的产生与清除的影响。阐述以脯氨酸为代表的渗透调节物质对活性氧的直接清除作用,Ca2+、甜菜碱等对抗氧化酶活性及抗氧化剂含量的影响。近年来人们广泛利用转基因技术合成脯氨酸、甜菜碱,为提高作物的抗氧化能力及培育抗逆新品种提供了一条有效途径。     

不同抗旱品种高粱苗中脯氨酸累积的差异

高粱抗旱品种“3197 B”比不抗旱品种“三尺三”在同样渗透胁迫条件下,要累积更多的游离脯氨酸。未受渗透胁迫前,两品种地上部的总氨基氮和游离氨基氮水平相似。氨基酸组成也相同。六种主要氨基酸分别占总的量和游离氨基酸量的50％和65％左右。受渗透胁迫24h后,总氨基氮水平未变,游离氨基氮增加,两品种的变化相似。因而,品种间脯氨酸累积的差异,似乎不因蛋白质或游离氨基酸的氨基酸组分的不同和蛋白质水解或合成受促进或抑制程度的不同而异。受渗透胁迫后,3197B植株内不但脯氨酸绝对量,而且相对量都超过“三尺三”。脯氨酸合成的潜在能力在3197B和“三尺三”之间有差别。     

渗透胁迫下稻苗中游离脯氨酸累积与膜脂过氧化的关系

杂交稻幼苗经聚乙二醇（ＰＧＥ４０００）渗透胁迫（－０．９５ＭＰａ）处理,幼苗含水量及相对含水量下降,游离脯氨酸和膜脂过氧化产物丙二醛（ＭＤＡ）含量上升,质膜透性增大。随ＰＥＧ渗透胁迫时间延长,幼苗膜脂饱和脂肪酸含量逐渐增加,不饱和脂肪酸含量降低,不饱和脂肪酸指数（ＩＵＦＡ）减少。脯氨酸累积与ＭＤＡ增长及膜透性加大呈正相关性,与膜脂脂肪酸不饱和度呈负相关性。讨论了游离脯氨酸累积与细胞透性的相关性,以     

干旱条件下植物ABA积累对脯氨酸水平的影响

目前公认,植物遭受水分胁迫后,体内发生适应性变化,最显著的是脱落酸(ABA)和脯氨酸的积累。ABA,能引起气孔关闭,调节植物体内水分平衡,保护质膜结构和功能,提高植物抗旱能力。脯氨酸作为渗透调     

Proteomic analysis of rice leaves shows the different regulations to osmotic stress and stress signals

Following the idea of partial root-zone drying(PRD)in crop cultivation,the morphological and physiological responses to partial root osmotic stress(PROS)and whole root osmotic stress(WROS)were investigated in rice.WROS caused stress symptoms like leaf rolling and membrane leakage.PROS stimulated stress signals,but did not cause severe leaf damage.By proteomic analysis,a total of 58 proteins showed differential expression after one or both treatments,and functional classification of these proteins suggests that stress signals regulate photosynthesis,carbohydrate and energy metabolism.Two other proteins(anthranilate synthase and submergence-induced nickel-binding protein)were upregulated only in the PROS plants,indicating their important roles in stress resistance.Additionally,more enzymes were involved in stress defense,redox homeostasis,lignin and ethylene synthesis in WROS leaves,suggesting a more comprehensive regulatory mechanism induced by osmotic stress.This study provides new insights into the complex molecular networks within plant leaves involved in the adaptation to osmotic stress and stress signals.     

Genotype differences in the metabolism of proline and polyamines under moderate drought in tomato plants

Water stress is one of the most important factors limiting the growth and productivity of crops. The implication of compatible osmolytes such as proline and polyamines in osmotic adjustment has been widely described in numerous plants species under stress conditions. In the present study, we investigated the response of five cherry tomato cultivars (Solanum lycopersicum L.) subjected to moderate water stress in order to shed light on the involvement of proline and polyamine metabolism in the mechanisms of tolerance to moderate water stress. Our results indicate that the most water stress‐resistant cultivar (Zarina) had increased degradation of proline associated with increased polyamine synthesis, with a higher concentration of spermidine and spermine under stress conditions. In contrast, Josefina, the cultivar most sensitive to water stress, showed a proline accumulation associated with increased synthesis after being subjected to stress. In turn, in this cultivar, no rise in polyamine synthesis was detected. Therefore, all the data appear to indicate that polyamine metabolism is more involved in the tolerance response to moderate water stress.     

Emerging physiological roles for N-acylphosphatidylethanolamine metabolism in plants: signal transduction and membrane protection

The activation of N-acylphosphatidylethanolamine (NAPE) metabolism in plants appears to be associated mostly with cellular stresses. In response to pathogen elicitors, NAPE is hydrolzyed by phospholipase-D (PLD), and corresponding medium-chain, saturated N-acylethanolamines (NAEs) are released by plant cells where they act as lipid mediators to modulate ion flux and activate defense gene expression. In desiccated seeds of higher plants, long-chain, saturated and unsaturated NAEs are prevalent, but are rapidly metabolized during the first few hours of imbibition, a period of substantial osmotic stress. NAPE synthesis is increased in seeds during this same period of rapid rehydration. A membrane-bound enzyme designated NAPE synthase has been purified from imbibed cottonseeds and its unusual biochemical properties suggest that it may scavenge free fatty acids in vivo. This feature of NAPE metabolism may be unique to higher plants a may be a mechanism for the rapid recycling of fatty acids back into membrane-associated NAPE. Altogether, increasing evidence indicates that NAPE metabolism in plants shares functional similarities with NAPE metabolism in animal systems, including signal transduction and cellular protection. In particular, the emerging role of released NAEs as lipid mediators in plant defense signaling represents an intriguing parallel to 'endocannabinoid signaling' in several mammalian cell types.     

Relationships between polyamines, ethylene, osmoprotectants and antioxidant enzymes activities in wheat seedlings after short-term PEG- and NaCl-induced stresses

Contents of ethylene, osmoprotectants, levels and forms of polyamines (PAs) and activities of antioxidant enzymes in the leaves and roots were investigated for five wheat cultivar seedlings (differing in drought tolerance) exposed to osmotic stress (?1.5 MPa). Stress was induced by 2-day-long treatment of plants with polyethylene glycol 6000 (PEG) or NaCl added to hydroponic cultures. Nawra, Parabola and Manu cv. (drought tolerant) showed a marked increase in osmoprotectors (proline and soluble carbohydrates, mainly glucose, saccharose and maltose), free PAs (putrescine Put, spermidine Spd and spermine Spm) and Spd-conjugated levels, in both leaves and roots, after PEG-treatments. Radunia and Raweta (drought sensitive) exhibited smaller changes in the content of these substances. The analysis of enzymes involved in proline metabolism revealed the glutamate as a precursor of proline synthesis in PEG-induced stress conditions. The increase in the activity of antioxidative enzymes, especially catalase and peroxidases, was characteristic for tolerant wheat plants, but for sensitive ones, a decrease in superoxide dismutase and an increase in mainly glutathione reductase activities were observed. After NaCl-treatment smaller changes of all biochemical parameters were registered in comparison with PEG-induced stress. Exceptions were the higher values of ethylene content and a significant increase in saccharose, raffinose and maltose levels (only in stress sensitive plants). The proline synthesis pathway was stimulated from both glutamate and ornithine precursors. These results suggest that the accumulation of inorganic ions in NaCl-stressed plants may be involved in protective mechanisms as an additional osmoregultor. Thus, a weaker stressogenic effect as determined as water deficit by leaf relative water content and relative dry weight increase rate and differences in metabolite synthesis in comparison with PEG stress was observed. Proline seems to be the most important osmo-protector in osmotic stress initiated by both PEG and NaCl. The synthesis of sugars and PAs may be stimulated in a stronger stress conditions (PEG).     

Effect of osmotic stress and sodium nitroprusside pretreatment on proline metabolism of wheat seedlings

Effect of osmotic stress and sodium nitroprusside (SNP, NO donor) pretreatment on growth and proline metabolism of wheat seedlings was investigated. Polyethylene glycol 6000 treatment for 2, 4 and 6 d could be termed as mild, moderate and severe stress, respectively, according to decrease in the relative water content. Severe osmotic stress significantly decreased the growth and photochemical efficiency, and increased proline content due to activation of its synthesis. 0.2 mM SNP pretreatment enhanced growth of wheat seedlings, increased variable to maximum fluorescence ratio (F_v/F_m) and fluorescence yield, while decreased proline content. However, 2 mM SNP retarded the seedlings growth and chlorophyll a fluorescence, and increased proline accumulation. Our results showed that NO might be involved in the regulation of osmotic stress in a concentration-dependent manner.     

植物甜菜碱合成酶的分子生物学和基因工程

甜菜碱是一种非毒性的渗透调节剂。多种高等植物在盐碱或缺水的环境下在细胞中积累甜菜碱 ,以维持细胞的正常膨压。甜菜碱的积累使得许多代谢中的重要酶类在渗透胁迫下能保持活性。在植物中甜菜碱由胆碱经两步氧化得到 ,催化第一步反应的酶是胆碱单加氧酶 (CMO) ,催化第二步反应的酶是甜菜碱醛脱氢酶 (BADH)。本文综述了这两种酶的分子生物学及基因工程研究的最新进展 ,讨论了其基因工程研究的意义。     

Expression of human dopamine receptor in potato (Solanum tuberosum) results in altered tuber carbon metabolism

Background  

Even though the catecholamines (dopamine, norepinephrine and epinephrine) have been detected in plants their role is poorly documented. Correlations between norepinephrine, soluble sugars and starch concentration have been recently reported for potato plants over-expressing tyrosine decarboxylase, the enzyme mediating the first step of catecholamine synthesis. More recently norepinephrine level was shown to significantly increase after osmotic stress, abscisic acid treatment and wounding. Therefore, it is possible that catecholamines might play a role in plant stress responses by modulating primary carbon metabolism, possibly by a mechanism similar to that in animal cells. Since to date no catecholamine receptor has been identified in plants we transformed potato plants with a cDNA encoding human dopamine receptor (HD1).     

Engineering of betaine biosynthesis and transport for abiotic stress tolerance in plants

Drought and salinity are the major factors that decrease crop yield. Organisms thriving in osmotic stress environments need adaptive mechanisms for adjusting their intracellular environment to external osmotic stress conditions. One such mechanism, to prevent water loss from the cells is to accumulate large amounts of low molecular weight organic compatible solutes such as proline, betaine and polyols to balance internal osmolarity of the cells. Accumulation of compatible solutes can be achieved by enhanced synthesis and/or reduced catabolism. Certain plants synthesize betaine in chloroplasts via a two-step oxidation of choline and betaine accumulation is associated with enhanced stress tolerance. Many important crop plants have low levels of betaine or none at all. Hence, betaine biosynthetic pathway is a target for metabolic engineering to enhance stress tolerance in crops. Introduction of betaine synthesis pathway into betaine non-accumulating plants has often improved stress tolerance. However, betaine levels of the engineered plants were generally low. To further enhance the betaine accumulation levels, we need to diagnose factors limitng betaine accumulation in engineered plants. Here we discuss recent progress on metabolic engineering of choline precursors for abiotic stress tolerance in plants.