水分胁迫下小麦类脱水素基因表达的半定量RT-PCR分析

以‘郑引1号’小麦为材料,经水分胁迫后进行RT-PCR扩增,结果在胁迫条件下获得500bp的脱水素基因(wzy1-1)。利用半定量RT-PCR方法,分析‘陕合6号’和‘郑引1号’小麦在正常供水条件下及PEG6000胁迫后18、24和42h,以及复水6和12h时wzy1-1在叶片中的表达。结果表明:2个小麦品种中wzy1-1在水分胁迫后18h均有表达,至胁迫24、48h时表达量较18h均有所增多;复水6h后该基因表达迅速降低,至复水后12h表达消失。且该基因在‘陕合6号’(抗旱性较强)叶片中表达量较‘郑引1号’(抗旱性较弱)高,表明该基因的表达与小麦的抗旱性密切相关。     

干旱胁迫下外源茉莉酸甲酯对玉米幼苗根系吸水的影响

茉莉酸类化合物作为环境信号分子,不仅参与植物生长发育的调控,同时受到环境胁迫的诱导,参与植物对逆境胁迫的响应和防御。本研究以北方广泛种植的玉米品种‘郑单958’为材料,通过对根系外源施加茉莉酸甲酯的方式,探究干旱胁迫下茉莉酸甲酯对玉米幼苗抗旱性以及根系吸水的影响。结果表明,外源茉莉酸甲酯可提高玉米幼苗光合速率、蒸腾作用和气孔导度,增强抗氧化酶活性,降低H_2O_2和丙二醛的含量,从而缓解干旱胁迫对植株造成的损伤。通过测定根系水导、氯化汞处理的蒸腾速率的变化以及水通道蛋白的表达量,发现干旱胁迫下外源茉莉酸甲酯可增强根系水通道蛋白的表达,进而增强玉米幼苗的根系吸水能力,从而缓解干旱胁迫造成的叶片水分含量的下降和水势的降低,提高了玉米幼苗的抗旱性。     

水分胁迫对燕麦穗颖渗透调节和抗氧化能力的影响

以抗旱性不同的燕麦品种‘蒙燕1号’(抗旱性强)和‘坝莜3号’(水分敏感)为试验材料,采用盆栽方式研究了抽穗期和灌浆期水分胁迫对燕麦穗颖渗透调节和抗氧化能力的影响。结果表明:(1)水分胁迫处理均显著促进了不同抗旱性品种穗颖渗透调节物质(游离脯氨酸和可溶性蛋白)含量增加,并以抗旱品种累积水平高于水敏感品种,且两种渗透调节物质对抽穗期胁迫的反应比灌浆期胁迫更敏感。(2)两时期的水分胁迫处理均能降低不同抗旱性品种穗颖SOD和POD活性,抗旱品种的保护酶活性要高于水敏感品种,抗旱品种的SOD活性降低幅度明显低于水敏感品种,而POD活性降低幅度在两品种间差异不明显。(3)水分胁迫导致2个品种穗颖丙二醛(MDA)含量和相对电导率显著增加,细胞膜结构受到严重伤害,且水敏感品种受害程度大于抗旱品种。(4)水分胁迫使2个品种单株籽粒产量下降,且在中度胁迫和重度胁迫下,抗旱品种的减产幅度要低于同期水敏感品种;水分胁迫下,水敏感品种‘坝莜3号’减产4.54%~30.29%,抗旱品种‘蒙燕1号’减产6.69%~23.54%。可见,抗旱性强的燕麦品种在受到水分胁迫的条件下能通过增强穗颖渗透调节和抗膜质过氧化能力、减弱穗颖细胞质膜损伤程度来适应干旱胁迫,最大限度减少水分胁迫对穗颖的伤害,有利于稳产。     

水分亏缺条件下玉米根系PIP2-5基因的表达(简报)

在PEG-6000模拟水分亏缺条件下,以微管蛋白基因为内参基因,水通道蛋白基因PIP2—5为检测基因,采用半定量逆转录聚合酶链式反应（RT-PCR）体系检测PIP2—5基因在玉米根系中的表达的结果表明,人工模拟水分亏缺条下PIP2—5基因表达量高于正常水分条件下的。这暗示,水分亏缺条件下细胞-细胞途径对根系吸水的贡献可能增大。     

沼液施用对干旱胁迫下玉米幼苗生长和生理特性的影响*

为了探明干旱胁迫下沼液对玉米幼苗抗旱、光合生理、形态的缓解效应,以中度抗旱玉米杂交种‘先玉335’和较强抗旱杂交种‘中单2号’为材料,采用10%聚乙二醇-6000模拟干旱胁迫,研究50%沼液根部浇灌处理对干旱胁迫下玉米幼苗生长和生理特性的影响。结果表明,沼液根部施用可以显著提高两品种玉米的抗旱性,有效缓解干旱胁迫对玉米幼苗根系和地上部生长的抑制作用,促进两品种玉米幼苗生长和提高根系活力,降低根冠比,且‘先玉335’的变幅更大;同时显著提高两品种叶片SOD、POD和CAT活性以及可溶性糖、脯氨酸、可溶性蛋白含量(‘中单2号’的CAT除外),降低MAD含量,且对‘中单2号’的影响更显著;沼液根施还可以显著提高干旱胁迫下两品种叶片的净光合速率(Pn)、蒸腾速率(Tr),降低气孔导度(Gs)、胞间CO_(2)浓度(Ci),但‘中单2号’的Tr和Gs除外,同时使两品种叶片叶绿素含量和水分利用效率(WUE)均显著增加。可见,沼液根施处理可以有效改善干旱胁迫下玉米幼苗的光合能力,显著提高幼苗抗氧化酶活性和渗透调节物质的含量,减轻膜脂过氧化程度,有效缓解干旱胁迫对2种不同抗旱性玉米幼苗的生长抑制,从而增强玉米耐受干旱胁迫的能力,且对‘中单2号’的缓解效果更明显。     

PEG胁迫及复水对不同抗旱性小麦幼苗脯氨酸代谢关键酶活性的影响

以两种不同抗旱性小麦品种幼苗为试验材料,采用PEG模拟干旱胁迫处理,探究干旱胁迫及复水对小麦幼苗叶片与根系脯氨酸累积及关键酶活性的影响。结果显示:(1)PEG胁迫下抗旱品种‘普冰143’根长和根干重下降不大,而水敏感品种‘郑引1号’根长和根干重下降显著;且于胁迫处理36h时‘普冰143’根系脯氨酸含量增加(75.0%)显著大于‘郑引1号’(37.7%),复水24h后均恢复至对照水平。(2)PEG胁迫下‘普冰143’叶片中谷氨酸合成途径关键酶P5CS和鸟氨酸合成途径关键酶δ-OAT活性均显著增加,且‘普冰143’叶片脯氨酸两条合成途径关键酶活性均得以加强;PEG胁迫处理36h时,‘郑引1号’叶片中P5CS活性增加显著,δ-OAT活性变化较小,且‘郑引1号’叶片脯氨酸合成可能以谷氨酸途径为主;但在PEG胁迫下两个不同抗旱性品种的根中P5CS、δ-OAT活性均变化较小。(3)PEG胁迫处理36h时‘普冰143’叶片脯氨酸降解酶PDH活性显著下降,而‘郑引1号’叶片PDH活性显著增加,复水后抗旱品种叶片该酶活性显著增加,水敏感品种恢复至对照水平;但PEG胁迫处理下两个不同抗旱性品种的根中PDH活性均显著下降。研究表明,PEG胁迫下小麦叶片是合成脯氨酸的主要部位,抗旱品种‘普冰143’根系脯氨酸持续积累与叶片中高的脯氨酸合成关键酶活性及脯氨酸转运有关。     

干旱胁迫及复水对不同黍稷品种根系生理特性的影响

以2种抗旱性不同的黍稷品种(‘陇糜4号’和‘晋黍7号’)为试验材料,采用盆栽试验研究了苗期中度和重度干旱胁迫后拔节期复水对其根系生理特性的影响。结果显示:(1)干旱胁迫引起2个黍稷品种根系活力明显下降,根系SOD、POD活性以及MDA、脯氨酸含量明显升高,而且重度干旱胁迫处理变化幅度显著大于中度干旱胁迫。(2)复水后,2个黍稷品种根系的各项生理指标均有不同程度的恢复,且中度胁迫处理较易恢复,重度胁迫下恢复能力很弱。(3)2个黍稷品种根系各项生理指标在干旱胁迫及复水条件下变化幅度不同,干旱胁迫下抗旱性强的‘陇糜4号’根系活力下降幅度明显低于抗旱性弱的‘晋黍7号’,根系SOD活性、POD活性、MDA含量和脯氨酸含量的上升幅度明显高于‘晋黍7号’,而复水后‘陇糜4号’根系的各项生理指标的恢复能力明显强于‘晋黍7号’。研究表明,干旱胁迫及复水条件下‘陇糜4号’均表现出较高的根系活力、保护酶活性和脯氨酸含量,且MDA含量较低,从而表现出较强的抗旱性。     

水分胁迫下花生叶片AhNCED1蛋白表达与气孔开度的变化

研究了水分胁迫下不同花生抗旱品种叶片气孔开度和相对含水量变化,分析TAhNCEDI基因和AhNCED1蛋白进行表达情况,发现水分胁迫下,叶片相对含水量下降,叶片气孔开度降低,叶片AhNCED1基因和AhNCED1蛋白表达增强。抗旱品种较之敏旱品种在响应水分胁迫初期时（1h）AhNCED1基因和AhNCED1蛋白表达较强,叶片气孔开度下降较快,引发气孔关闭,其叶片相对含水量较高,保水能力较强。ABA合成抑制剂naproxen处理后,叶片AhNCED1基因和AhNCED1蛋白的表达减弱,气孔开度快速增加,水分胁迫下花生叶片AhNCED1蛋白表达可能影响气孔开闭。     

干旱胁迫对不同苦荞品种苗期生长和根系生理特征的影响

采用盆栽人工控水试验,研究了不同水分处理(正常供水、中度干旱和重度干旱)对耐旱型(‘迪庆苦荞’、‘西农9909’)和不耐旱型(‘西荞1号’和‘黑丰1号’)苦荞品种苗期生理、形态指标的影响,并通过隶属函数法与主成分分析对品种抗旱性进行综合评价,以揭示苦荞苗期的抗旱生理机制。结果表明:(1)与正常供水相比,除‘迪庆苦荞’和‘西农9909’在重度干旱胁迫下主根长呈升高趋势外,其余苦荞品种在2个干旱条件下的株高、茎粗、叶面积、地上部干重、地下部干重、根系体积、根系表面积均呈下降趋势,且耐旱品种降幅小于不耐旱品种;重度干旱胁迫使得‘迪庆苦荞’的根冠比升高,而其余品种根冠比在干旱胁迫下均无显著变化。(2)干旱胁迫使苦荞叶片的叶绿素含量、相对含水量、最大荧光产量(Fm)、最大光化学效率(Fv/Fm)、根系活力和可溶性蛋白含量显著降低,而根系超氧化物歧化酶(SOD)活性、过氧化物酶(POD)活性、丙二醛(MDA)、可溶性糖及游离脯氨酸含量呈升高趋势;不同抗旱性品种间的升降幅度存在差异。(3)各苦荞品种耐旱能力综合评价值(D)表现为‘迪庆苦荞’‘西农9909’‘黑丰1号’‘西荞1号’;幼苗株高、地下部干重及根系SOD活性和蛋白质含量与D值呈显著正相关关系,而根系脯氨酸含量和可溶性糖含量与D值呈极显著正相关关系。研究发现,在中度与重度干旱逆境下,苦荞品种‘迪庆苦荞’和‘西农9909’综合表现较好,具有更强的耐旱能力,而品种‘西荞1号’和‘黑丰1号’综合表现较差,其抗旱性较弱;苗期株高、地下部干重以及根系SOD活性、蛋白质含量、脯氨酸含量和可溶性糖含量可作为苦荞抗旱性快速鉴定的指标。     

水分胁迫对不同基因型小麦幼芽蛋白质表达和某些生理特性的影响

以抗旱小麦品种长武134和不抗旱小麦品种郑引1号为试材,采用-1.2 MPa PEG 6000处理种子,研究不同水分条件下小麦幼芽中蛋白质表达和部分生理特性的变化.聚丙烯酰胺凝胶电泳结果表明：水分胁迫可诱导抗旱品种幼芽产生分子量约39.5 kDa和23.0 kDa两种新蛋白亚基,不抗旱品种则无新亚基产生;在正常供水条件下,随着生育期延长,两品种中分子量为48.5 kDa的蛋白亚基表达量逐渐增强,因其对水分敏感且属于新发现蛋白,初步命名为水敏感蛋白.生理特性测定结果表明,水分胁迫条件下长武134的根芽比和相对含水量均高于郑引1号,而细胞膜相对透性和丙二醛含量则低于郑引1号.     

Short-term control of maize cell and root water permeability through plasma membrane aquaporin isoforms

Although it is widely accepted that aquaporins are involved in the regulation of root water uptake, the role of specific isoforms in this process is poorly understood. The mRNA expression and protein level of specific plasma membrane intrinsic proteins (PIPs) were analysed in Zea mays in relation to cell and root hydraulic conductivity. Plants were analysed during the day/night period, under different growth conditions (aeroponics/hydroponics) and in response to short-term osmotic stress applied through polyethylene glycol (PEG). Higher protein levels of ZmPIP1;2, ZmPIP2;1/2;2, ZmPIP2;5 and ZmPIP2;6 during the day coincided with a higher water permeability of root cortex cells during the day compared with night period. Similarly, plants which were grown under aeroponic conditions and which developed a hypodermis ('exodermis') with Casparian bands, effectively forcing more water along a membranous uptake path across roots, showed increased levels of ZmPIP2;5 and ZmPIP1;2 in the rhizodermis and exodermis. When PEG was added to the root medium (2-8 h), expression of PIPs and cell water permeability in roots increased. These data support a role of specific PIP isoforms, in particular ZmPIP1;2 and ZmPIP2;5, in regulating root water uptake and cortex cell hydraulic conductivity in maize.     

Coronatine promotes maize water uptake by directly binding to the aquaporin ZmPIP2;5 and enhancing its activity

Water uptake is crucial for crop growth and development and drought stress tolerance. The water channel aquaporins (AQP) play important roles in plant water uptake. Here, we discovered that a jasmonic acid analog, coronatine (COR), enhanced maize (Zea mays) root water uptake capacity under artificial water deficiency conditions. COR treatment induced the expression of the AQP gene Plasma membrane intrinsic protein 2;5 (ZmPIP2;5). In vivo and in vitro experiments indicated that COR also directly acts on ZmPIP2;5 to improve water uptake in maize and Xenopus oocytes. The leaf water potential and hydraulic conductivity of roots growing under hyperosmotic conditions were higher in ZmPIP2;5-overexpression lines and lower in the zmpip2;5 knockout mutant, compared to wild-type plants. Based on a comparison between ZmPIP2;5 and other PIP2s, we predicted that COR may bind to the functional site in loop E of ZmPIP2;5. We confirmed this prediction by surface plasmon resonance technology and a microscale thermophoresis assay, and showed that deleting the binding motif greatly reduced COR binding. We identified the N241 residue as the COR-specific binding site, which may activate the channel of the AQP tetramer and increase water transport activity, which may facilitate water uptake under hyperosmotic stress.     

Differential responses of maize MIP genes to salt stress and ABA

Salt stress is known to reduce root hydraulic conductivity and growth. To examine a concomitant regulation of aquaporins, the expression of the maize MIP gene family in response to NaCl was analysed by DNA array hybridization. Plants responded differentially to 100 versus 200 mM NaCl treatments. Leaf water content was reduced rapidly and persistently after the application of 200 mM NaCl in contrast to 100 mM NaCl. Endogenous ABA strongly accumulated in roots after 2 h; it remained at a highly elevated level for 48 h after the addition of 200 mM NaCl, but rapidly declined in plants treated with 100 mM NaCl, indicating an early recovery from water deficit. Interestingly, 2 h after the addition of 100 mM NaCl, when maize regained the osmotic potential allowing water uptake, three highly expressed, specific isoforms ZmPIP1;1, ZmPIP1;5, and ZmPIP2;4 were transiently induced. They were preferentially transcribed in the outer root tissue suggesting a role in cellular water transport. None of the ZmTIP genes was altered. By contrast, after the addition of 200 mM NaCl these responses were missing. Instead, multiple ZmPIP and ZmTIP genes were repressed by 200 mM NaCl after 24 h. After 48 h, deregulations were overridden in both cases indicating homeostasis. ABA (1 muM) exogenously applied to the roots transiently induced ZmPIP2;4 similar to 100 mM NaCl as well as ZmPIP1;2. Thus, the early induction of ZmPIP2;4 by NaCl may be mediated by ABA. Previously, an increase in root hydraulic conductivity had been observed upon ABA application. By contrast, 100 muM ABA led to a complete, possibly non-specific repression of all detected ZmPIP and ZmTIP genes after 24 h.     

Aquaporin JcPIP2 is involved in drought responses in Jatropha curcas

Water channel proteins, aquaporins, play fundamental roles in transmembrane water movements in plants. A new full-length cDNA encoding aquaporin was isolated from the seedlings of Jatropha curcas. The gene of the plasma membrane intrinsic protein （PIP） from J. curcas （JcPIP2） contained an 843 bp open reading frame encoding a protein of 280 amino acids. The amino acid sequence showed 94% identity with Ricinus communis PIP. Injection of JcPIP2 complementary RNA into Xenopus oocytes increased 10-fold the osmotic water permeability of the oocytes. Immunodetection of JcPIP2 with anti-JcPIP2 antibody indicated that this protein is ubiquitously located in all tested tissues of the plant. To investigate the relationship between aquaporins and drought resistance in J. curcas, the abundance of JcPIP2 was examined in seedlings of two J. curcas populations, GaoYou CSC63 and YanBian S 1, under water deficit with PEG6000. Under field conditions, those two populations, GaoYou CSC63 was resistant to water deficit, but YanBian S 1 was sensitive to water deprivation. With the increasing degree of drought stress, JcPIP2 level increased in seedlings of GaoYou CSC63, whereas there was no significant change in seedlings of YanBian S 1. Compared with YanBian S 1, GaoYou CSC63 also showed higher root hydraulic conductivity and lower decreasing trend in the seed- lings under water deficit. These results indicated that JcPIP2 probably played a role in drought resistance in J. curcas.     

Plasma membrane intrinsic proteins from maize cluster in two sequence subgroups with differential aquaporin activity

The transport of water through membranes is regulated in part by aquaporins or water channel proteins. These proteins are members of the larger family of major intrinsic proteins (MIPs). Plant aquaporins are categorized as either tonoplast intrinsic proteins (TIPs) or plasma membrane intrinsic proteins (PIPs). Sequence analysis shows that PIPs form several subclasses. We report on the characterization of three maize (Zea mays) PIPs belonging to the PIP1 and PIP2 subfamilies (ZmPIP1a, ZmPIP1b, and ZmPIP2a). The ZmPIP2a clone has normal aquaporin activity in Xenopus laevis oocytes. ZmPIP1a and ZmPIP1b have no activity, and a review of the literature shows that most PIP1 proteins identified in other plants have no or very low activity in oocytes. Arabidopsis PIP1 proteins are the only exception. Control experiments show that this lack of activity of maize PIP1 proteins is not caused by their failure to arrive at the plasma membrane of the oocytes. ZmPIP1b also does not appear to facilitate the transport of any of the small solutes tried (glycerol, choline, ethanol, urea, and amino acids). These results are discussed in relationship to the function and regulation of the PIP family of aquaporins.     

玉米根系水流导度差异及其与解剖结构的关系

在人工气候室水培条件下,从单根水平研究了不同水分条件下玉米根系水流导度的基因型差异及解剖结构之间的关系.结果表明,抗旱性的杂交种户单四号具有水流导度上的杂种优势现象,不抗旱的父本803根系水流导度最低,3个品种根系水流导度大小为F₁代户单四号＞母本天四＞父本803;水分胁迫普遍降低了根系直径、导管直径和皮层厚度.同时,玉米品种根系的解剖结构和根系水流导度有关,正常水分条件下,根系导管直径与3个玉米品种的根系水流导度呈正相关,胁迫条件下则呈负相关.无论是在胁迫还是正常水分条件下,根系皮层厚度占根系直径的比例与根系水流导度都呈负相关,说明根系皮层是根系吸收水分的主要阻力部位.     

Localization and Quantification of Plasma Membrane Aquaporin Expression in Maize Primary Root: A Clue to Understanding their Role as Cellular Plumbers

Water movement across root tissues occurs by parallel apoplastic, symplastic, and transcellular pathways that the plant can control to a certain extent. Because water channels or aquaporins (AQPs) play an important role in regulating water flow, studies on AQP mRNA and protein expression in different root tissues are essential. Here, we quantified and localized the expression of Zea mays plasma membrane AQPs (ZmPIPs) in primary root tip using in situ and quantitative RT-PCR and immunodetection approaches. All ZmPIP genes except ZmPIP2;7 were expressed in primary roots. Expression was found to be dependent on the developmental stage of the root, with, in general, an increase in expression towards the elongation and mature zones. Two genes, ZmPIP1;5 and ZmPIP2;5, showed the greatest increase in expression (up to 11- and 17-fold, respectively) in the mature zone, where they accounted for 50% of the total expressed ZmPIPs. The immunocytochemical localization of ZmPIP2;1 and ZmPIP2;5 in the exodermis and endodermis indicated that they are involved in root radial water movement. In addition, we detected a polar localization of ZmPIP2;5 to the external periclinal side of epidermal cells in root apices, suggesting an important role in water uptake from the root surface. Finally, protoplast swelling assays showed that root cells display a variable, but globally low, osmotic water permeability coefficient (P _f < 10 μm/s). However, the presence of a population of cells with a higher P _f (up to 26 μm/s) in mature zone of the root might be correlated with the increased expression of several ZmPIP genes.     

An N-terminal diacidic motif is required for the trafficking of maize aquaporins ZmPIP2;4 and ZmPIP2;5 to the plasma membrane

Maize plasma membrane aquaporins (ZmPIPs, where PIP is the plasma membrane intrinsic protein) fall into two groups, ZmPIP1s and ZmPIP2s, which, when expressed alone in mesophyll protoplasts, are found in different subcellular locations. Whereas ZmPIP1s are retained in the endoplasmic reticulum (ER), ZmPIP2s are found in the plasma membrane (PM). We previously showed that, when co-expressed with ZmPIP2s, ZmPIP1s are relocalized to the PM, and that this relocalization results from the formation of hetero-oligomers between ZmPIP1s and ZmPIP2s. To determine the domains responsible for the ER retention and PM localization, respectively, of ZmPIP1s and ZmPIP2s, truncated and mutated ZmPIPs were generated, together with chimeric proteins created by swapping the N- or C-terminal regions of ZmPIP2s and ZmPIP1s. These mutated proteins were fused to the mYFP and/or mCFP, and the fusion proteins were expressed in maize mesophyll protoplasts, and were then localized by microscopy. This allowed us to identify a diacidic motif, DIE (Asp-Ile-Glu), at position 4–6 of the N-terminus of ZmPIP2;5, that is essential for ER export. This motif was conserved and functional in ZmPIP2;4, but was absent in ZmPIP2;1. In addition, we showed that the N-terminus of ZmPIP2;5 was not sufficient to cause the export of ZmPIP1;2 from the ER. A study of ZmPIP1;2 mutants suggested that the N- and C-termini of this protein are probably not involved in ER retention. Together, these results show that the trafficking of maize PM aquaporins is differentially regulated depending on the isoform, and involves a specific signal and mechanism.     

Boric acid and salinity effects on maize roots. Response of aquaporins ZmPIP1 and ZmPIP2, and plasma membrane H+-ATPase, in relation to water and nutrient uptake

Under saline conditions, an optimal cell water balance, possibly mediated by aquaporins, is important to maintain the whole-plant water status. Furthermore, excessive accumulation of boric acid in the soil solution can be observed in saline soils. In this work, the interaction between salinity and excess boron with respect to the root hydraulic conductance (L₀), abundance of aquaporins (ZmPIP1 and ZmPIP2), ATPase activity and root sap nutrient content, in the highly boron- and salt-tolerant Zea mays L. cv. amylacea, was evaluated. A downregulation of root ZmPIP1 and ZmPIP2 aquaporin contents were observed in NaCl-treated plants in agreement with the L₀ measurements. However, in the H₃BO₃-treated plants differences in the ZmPIP1 and ZmPIP2 abundance were observed. The ATPase activity was related directly to the amount of ATPase protein and Na⁺ concentration in the roots, for which an increase in NaCl- and H₃BO₃+ NaCl-treated plants was observed with respect to untreated and H₃BO₃-treated plants. Although nutrient imbalance may result from the effect of salinity or H₃BO₃ alone, an ameliorative effect was observed when both treatments were applied together. In conclusion, our results suggest that under salt stress, the activity of specific membrane components can be influenced directly by boric acid, regulating the functions of certain aquaporin isoforms and ATPase as possible components of the salinity tolerance mechanism.