植物质膜钾离子转运体研究进展

近年,随着分子生物学技术的不断发展和广泛应用,有关植物质膜钾离子转运体的研究取得重要进展。目前已经克隆到多种质膜钾离子转运体基因并对钾离子转运体生化特性以及结构功能进行了广泛研究。研究认为,质膜钾离子转运体可分为钾离子载体和钾离子通道。钾离子通道又可分为内向性K+通道α亚基、K+通道β亚基及外向性K+通道等三类。本文对上述质膜钾离子转运体的生化特性以及结构功能研究的进展进行了综述。     

植物质膜钾离子转运体研究进展

近年,随着分子生物学技术的不断发展和广泛应用,有关植物质膜钾离子转运体的研究取得重要进展。目前已经克隆到多种质膜钾离子转运体基因并对钾离子转运体生化特性以及结构功能进行广泛研究。研究认为,质膜钾离子转运体可分为钾离子载体和钾离子通道。钾离子通道又可分为内向性Ｋ＾＋通道α亚基、Ｋ＾＋通道β亚基及外生Ｋ＾＋通道等三类。本文对上述质膜钾离子转运体的生化特性以及结构功能研究的进展进行了综述。     

硅缓解植物镉毒害的生理生态机制

镉是对生物毒性最强的污染物之一。过量的镉能够抑制植物的生长和光合作用,干扰矿质代谢并诱发氧化胁迫。硅作为一种有益元素,主要以Si(OH)_4的形态通过主动或被动方式被植物体吸收并转运到地上部分。硅对植物镉毒害具有缓解作用,但其缓解机制在不同物种、品种或生态型之间存在显著差异,并表现出一定的硅/镉浓度依赖性。总体上可概括为避性机制和耐性机制。避性机制包括:(1)在器官水平,减少植物根系对镉的吸收及其向地上部的转运;(2)在细胞水平,增强细胞壁对镉的吸附能力,减少共质体中镉的含量。耐性机制包括:(1)诱导细胞产生小分子螯合剂,增强对镉的螯合作用,减少细胞中游离态镉的含量;(2)增强抗氧化机制,减轻氧化胁迫;(3)改善光合作用和无机营养,促进植物生长。从植物对硅的吸收和转运、镉对植物的毒害作用以及硅对缓解植物镉毒害的生理生态机制3个方面进行了综述,并基于目前的研究现状和薄弱之处,对今后的研究重点进行了展望。     

根表铁氧化物胶膜对水稻吸收Zn的影响

采用营养液培养方法研究了水稻根表形成的铁氧化物胶膜对水稻吸收Ｚｎ的影响．结果表明,在有Ｆｅ２＋的嫌气环境中,由于根际氧化作用水稻根表会形成红色的铁氧化物胶膜,根表的铁氧化物胶膜影响水稻对Ｚｎ的吸收．铁膜数量较少时,由于对Ｚｎ的富集作用有限,其对水稻Ｚｎ的吸收虽有促进作用,但不明显．随着根表铁膜数量的增加,这种促进作用也相应增加,并且在铁膜数量增加到一定值时,对水稻吸收Ｚｎ的促进作用达到最大．而后,随着铁膜数量的进一步增加,铁膜反而阻碍水稻对Ｚｎ的吸收,成为水稻吸收Ｚｎ的障碍层．在此过程中,水稻的根分泌物,特别是其中的植物铁载体对覆有铁膜水稻根系吸收Ｚｎ有促进作用．这种促进作用随铁膜数量的增加而逐渐减弱．因此,根表铁氧化物胶膜对水稻吸收Ｚｎ并不总是起促进作用,其作用的方向和程度取决于铁膜的数量．     

秋茄对镉的吸收、积累及净化作用的研究

采用砂培研究红树植物秋茄幼苗在含镉0.0025—25ppm海水培养液中对镉的吸收积累、净化作用以及镉对秋茄的毒害作用。试验结果表明：秋茄植物体镉含量大小顺序为：根>胚轴>茎>叶．各器官镉含量(y)随培养液镉浓度(x)的增加而增加,两者呈显著的双对数线性正相关,根与培养液镉含量的回归方程为;logy=1.8039+0.4311logx。在含镉培养液中培养4个星期,不同器官对镉的富集作用明显,镉处理2.5ppb,叶、茎、胚轴、根的富集系数分别为62.4、113.6、187.2、705.2．吸收的镉主要积累于根部。25ppm镉处理组每株幼苗(干重6.18g)吸收镉量为239.47μg,其中根占植物体总镉量的75.6%．在所有浓度处理中,秋茄植物体对镉的吸收量为培养液总加镉量的2.4—36%。2.5ppm镉处理组10个星期内,根部镉积累量为538.74μg／g,未见明显受害,表明秋茄对镉的吸收积累能力和耐性都较强。幼苗受镉的毒害程度主要与根部镉含量有关。从天然条件生长的红树植物器官镉积累量来看,也表明秋茄群落对镉污染的吸收净化效果显著。     

不同营养环境条件下铜、锌、锰等二价金属营养元素对水稻吸收运转镉的调控

利用水培试验研究了在不同营养环境条件下几种二价金属营养元素与镉的互作关系及其水稻吸收运转镉的影响。研究结果表明,培养液中全部营养元素供应水平变化,二价金属元素浓度变化均对水稻吸收镉产生显著影响。Cd浓度保持为2μmol/L不变,随着全部必须元素浓度系数从0. 25提高到2. 0,水稻根系和茎叶Cd含量分别降低42. 5%和64. 5%。在没有其它二价阳离子干扰的情况下,Mg^(2+)、Fe^(2+)、Zn^(2+)、Mn^(2+)、Cu^(2+)均强烈抑制水稻对镉的吸收,水稻根系镉含量降低至对照的15. 4%～61. 6%,茎叶镉含量降低至对照的20. 2%～87. 5%。在木村B培养液中增量供应Zn^(2+)或Mn^(2+)均大幅度促进镉在根系、茎叶和稻谷积累,其中增量供应Zn^(2+)处理的水稻根、茎叶和稻谷镉含量分别比对照增加45. 5%、177. 4%、250. 8%,而增量供应Mn^(2+)处理的水稻根系、茎叶、稻谷镉含量分别比对照增加20. 2%、117. 4%、227. 1%。在所有二价金属元素中,Cu^(2+)对水稻吸收Cd^(2+)的拮抗作用最强,而且铜-镉拮抗关系受营养环境的影响较小。培养液中增量供应Cu^(2+)导致水稻吸收镉大幅减少,根系、茎叶、稻谷镉含量分别比对照降低51. 4%、45. 2%、52. 2%;培养液缺Cu^(2+)则导致根系和茎叶镉含量分别比对照增加177%和114%。研究结果表明,在镉污染稻田适度施用铜肥、控制锌锰的输入、在水稻生长发育期保持较高土壤营养水平可能是控制水稻积累镉的营养调控方法。     

植物根毛的发生、发育及养分吸收

根毛是植物吸收养分的重要器官,认识根毛的发生、发育规律及其与养分吸收的关系,可为植物养分吸收效率的遗传改良提供依据。介绍了植物根毛的形态特性、发生和发育过程及其调控机制,并结合本实验室的工作,讨论了根毛对养分吸收的贡献、根毛受养分有效性的调节及其与其他根系形态构型性状间的关系,阐述了根毛中养分转运等植物营养过程及其生理和分子生物学基础。最后提出了关于根毛研究中的一些问题和研究前景。     

植物根毛的发生、发育及养分吸收

根毛是植物吸收养分的重要器官,认识根毛的发生、发育规律及其与养分吸收的关系,可为植物养分吸收效率的遗传改良提供依据.介绍了植物根毛的形态特性、发生和发育过程及其调控机制,并结合本实验室的工作,讨论了根毛对养分吸收的贡献、根毛受养分有效性的调节及其与其他根系形态构型性状间的关系,阐述了根毛中养分转运等植物营养过程及其生理和分子生物学基础.最后提出了关于根毛研究中的一些问题和研究前景.     

AM真菌提高烟草钾含量机制的初步研究

以烟草和AM真菌菌株摩西球囊霉（Glomous mosseae,G.m）为材料,研究了不同外界K＋浓度条件下AM真菌对烟草生长、K＋含量以及根中K＋-通道基因NtKT1、NtKT2和转运体基因NtHAK1、NtHA1相对表达量的影响。结果表明,在低钾和常钾条件下,接种G.m均可提高烟草的株高,增加根系长度,提高烟草根部和叶片中的K＋含量,在低钾条件下接种G.m使烟草根部和叶片的K＋含量分别提高了50.5%和24.5%。在常钾条件下,接种G.m 50 d后烟草根系中NtKT1和NtKT2的相对表达量显著提高;而低钾条件下,钾转运体基因NtHAK和NtHA1的相对表达量显著升高。由此推测,AM真菌能够调控烟草根中K＋-通道基因和钾转运体基因的表达进而促进其对K＋的吸收。     

植物HKT蛋白耐盐机制研究进展

盐害是限制植物生长发育的重要环境因素, 对植物造成渗透胁迫和离子毒害。维持细胞及整株水平的Na⁺/K⁺平衡是植物重要的耐盐机制。目前, 已报道的高亲和性钾离子转运蛋白(HKT)具有钠、钾离子转运特性, 在植物体钠、钾离子长距离运输及分配过程中发挥重要作用。该文重点总结了淡土植物和盐土植物HKT蛋白的结构、功能及耐盐机理, 并对其在植物耐盐改良育种中的前景做出了展望。     

Cd-K双重处理对水稻幼苗生长和Cd吸收转运的影响及相关转运通道分析

以水稻( Oryza sativa Linn.)高 Cd 积累品种‘T 优705’(‘T You 705’)和低 Cd 积累品种‘湘早籼24’(‘Xiangzaoxian 24’)为实验材料,采用水培法对不同浓度Cd(0.0和2.7μmol·L-1 Cd)和K(0、30和60 mmol·L-1 K)处理条件下2个品种幼苗的相对生长量、根系和地上部的Cd含量及其亚细胞分布特征进行了比较,并分析了添加离子通道活性抑制剂TEA和LaCl3后幼苗根系和地上部的Cd和K含量;在此基础上,比较了NSCCs(非选择性阳离子通道)和K专性通道对2个品种幼苗根系和地上部Cd和K吸收贡献率的影响。结果表明：与Cd单一处理组(2.7μmol·L-1 Cd)相比, Cd-K双重处理组(2.7μmol·L-1 Cd-30 mmol·L-1 K和2.7μmol·L-1 Cd-60 mmol·L-1 K)2个品种幼苗的相对生长量显著提高,而幼苗根系和地上部的Cd含量显著下降;随K浓度的提高,2个品种幼苗根系细胞壁和细胞液中的Cd含量显著下降,但细胞壁中Cd含量的分配比例增大而细胞液中Cd含量的分配比例则减小。在含2.7μmol·L-1 Cd和30 mmol·L-1 K的培养液中分别添加5 mmol·L-1 TEA或0.2 mmol·L-1 LaCl3后,2个品种幼苗根系和地上部的Cd和K含量均显著下降,其中,LaCl3处理组的根系Cd含量降幅高于TEA处理组,但LaCl3处理组的根系K含量降幅则低于TEA组。 NSCCs对品种‘T优705’幼苗根系和地上部Cd吸收的贡献率显著低于品种‘湘早籼24’幼苗,而K专性通道对品种‘T优705’幼苗根系K吸收和地上部Cd吸收的贡献率则显著低于品种‘湘早籼24’幼苗。研究结果显示：添加外源K可缓解Cd对水稻幼苗生长的抑制作用,并通过提高细胞壁与Cd的结合能力来降低细胞液中Cd的积累,以此减弱幼苗对Cd的吸收和转运能力;幼苗体内的K和Cd均可通过K专性通道和NSCCs转运,其中,K吸收和转运主要通过K专性通道完成,而Cd吸收和转运主要通过NSCCs完成。此外,品种‘T优705’可能具有多种离子通道参与Cd的吸收和转运,而品种‘湘早籼24’主要依赖NSCCs参与Cd的吸收和转运,且后者对K的吸收和积累强于前者。     

N、P、K肥对香根草修复土壤镉、锌污染效率的影响

通过盆栽试验研究在30 mg/kg镉(Cd)污染土壤条件下N[CO(NH2)2:100、200、300 mg/kg土]、P(P2O5:50、100、200 mg/kg土)和K(KCl:100、200、300 mg/kg土)处理对香根草修复土壤Cd和锌(Zn)污染效率的影响。结果表明:3种N处理能促进香根草地上部生长,而且显著提高地上部特别是叶的Cd和Zn含量,导致其修复效率成倍显著增加;200 mg/kg K处理显著提高Zn修复效率,但300 mg/kg K和50、200 mg/kg P处理却显著降低Cd、Zn修复效率。因此,为改善香根草对较贫瘠土壤中Cd、Zn污染的修复效率,应对香根草适施N肥,并控制或者不施P、K肥为佳。     

Cd2+ regulation of the hyperpolarization-activated current IAB in crayfish muscle

The effects of Cd2+ on the hyperpolarization-activated K(+)-mediated current called IAB (Araque, A., and W. Buno. 1994. Journal of Neuroscience. 14:399-408.) were studied under two-electrode voltage- clamp in opener muscle fibers of the crayfish Procambarus clarkii. IAB was reversibly reduced by extracellular Cd2+ in a concentration- dependent manner, obeying the Hill equation with IC50 = 0.452 +/- 0.045 mM and a Hill coefficient of 1 (determined from the maximal chord conductance of IAB). Cd2+ decreased the IAB conductance (GAB) and shifted its voltage dependence towards hyperpolarized potentials in a similar degree, without affecting the slope of the voltage dependence. The IAB activation time constant increased, whereas the IAB deactivation time constant was not modified by Cd2+. The IAB equilibrium potential (EAB) was unmodified by Cd2+, indicating that the selective permeability of IAB channels was not altered. IAB was unaffected by intracellular Cd2+. The Cd(2+)-regulation of IAB did not depend on [K+]o, and the effects of [K+]o on IAB were unchanged by Cd2+, indicating that Cd2+ did not compete with K+. Therefore, Cd2+ probably bound to a different site to that involved in the K+ permeability pathway. We conclude that Cd2+ affected the gating of IAB channels, interfering with their opening but not with their closing mechanism. The results can be explained by a kinetic model in which the binding of Cd2+ to the IAB channels would stabilize the gating apparatus at its resting position, increasing the energy barrier for the transition from the closed to the open channel states.     

Accumulation,distribution and toxicological effects induced by cadmium on the development of mangrove plant <Emphasis Type="Italic">Kandelia candel</Emphasis> (L.)

Accumulation and distribution pattern of cadmium (Cd) and its toxic effect on growth of the mangrove plant of young Kandelia candel seedlings have been examined. This study demonstrated that under high concentration of Cd stress, the total biomass of K. candel decreased 41.57% compare to control (CK). At the end of 90 days exposure to 25 mg/l Cd, the average seedlings stem height and leaf number of the K. candel decreased by 30.54 and 42.68%, respectively. The results showed that K. candel seedlings, under the experimental condition, accumulated higher concentration of Cd in their roots (411.29 ± 3.60 mg/kg) when compared to hypocotyls, stems and leaves. More than 95% of Cd was accumulated mainly in roots. The distribution pattern of Cd concentration in K. candel seedlings was found in the following order: roots > hypocotyls > stems > leaves. Based on the leaf symptoms and morphological change of K. candel seedlings under heavy metal stress, this study showed that Cd is phytotoxic to K. candel.     

Enhancement of HERG K(+) currents by Cd(2+) destabilization of the inactivated state

We have studied the functional effects of extracellular Cd(2+) on human ether-a-go-go-related gene (HERG) encoded K(+) channels. Low concentrations (10-200 &mgr;M) of extracellular Cd(2+) increased outward currents through HERG channels; 200 &mgr;M Cd(2+) more than doubled HERG currents and altered current kinetics. Cd(2+) concentrations up to 200 &mgr;M did not change the voltage dependence of channel activation, but shifted the voltage dependence of inactivation to more depolarized membrane potentials. Cd(2+) concentrations >/=500 &mgr;M shifted the voltage dependence of channel activation to more positive potentials. These results are consistent with a somewhat specific ability of Cd(2+) to destabilize the inactivated state. We tested the hypothesis that channel inactivation is essential for Cd(2+)-induced increases in HERG K(+) currents, using a double point mutation (G628C/S631C) that diminishes HERG inactivation (Smith, P. L., T. Baukrowitz, and G. Yellen. 1996. Nature (Lond.). 379:833-836). This inactivation-removed mutant is insensitive to low concentrations of Cd(2+). Thus, Cd(2+) had two distinct effects on HERG K(+) channels. Low concentrations of Cd(2+) caused relatively selective effects on inactivation, resulting in a reduction of the apparent rectification of the channel and thereby increasing HERG K(+) currents. Higher Cd(2+) concentrations affected activation gating as well, possibly by a surface charge screening mechanism or by association with a lower affinity site.     

Effect of fertilizers on Cd uptake of Amaranthus hypochondriacus, a high biomass, fast growing and easily cultivated potential Cd hyperaccumulator

In a greenhouse pot experiment, we assessed the phytoextraction potential for Cd of three amaranth cultivars (Amaranthus hypochondriacus L. Cvs. K112, R104, and K472) and the effect of application of N, NP, and NPK fertilizer on Cd uptake of the three cultivars from soil contaminated with 5 mg kg(-1) Cd. All three amaranth cultivars had high levels of Cd concentration in their tissues, which ranged from 95.1 to 179.1 mg kg(-1) in leaves, 58.9 to 95.4 mg kg(-1) in stems, and 62.4 to 107.2 mg kg(-1) in roots, resulting in average bioaccumulation factors ranging from 17.7 to 29.7. Application of N, NP, or NPK fertilizers usually increased Cd content in leaves but decreased Cd content in stem and root. Fertilizers of N or NP combined did not substantially increase dry biomass of the 3 cultivars, leading to a limited increment of Cd accumulation. NPK fertilizer greatly increased dry biomass, by a factor of 2.7-3.8, resulting in a large increment of Cd accumulation. Amaranth cultivars (K112, R104, and K472) have great potential in phytoextraction of Cd contaminated soil. They have the merits of high Cd content in tissues, high biomass, easy cultivation and little effect on Cd uptake by fertilization.     

Cadmium-mediated oxidative stress in kidney proximal tubule cells induces degradation of Na+/K(+)-ATPase through proteasomal and endo-/lysosomal proteolytic pathways.

The mechanisms of cadmium (Cd)-dependent nephrotoxicity were studied in a rat proximal tubule (PT) cell line. CdCl(2) (5 microM) increased the production of reactive oxygen species (ROS), as determined by oxidation of dihydrorhodamine 123 to fluorescent rhodamine 123. The levels of ubiquitin-conjugated cellular proteins were increased by Cd in a time-dependent fashion (maximum at 24-48 h). This was prevented by coincubation with the thiol antioxidant N-acetylcysteine (NAC, 15 mM). Cd also increased apoptosis (controls: 2.4+/-1.6%; Cd: 8.1+/-1.9%), but not necrosis (controls: 0.5 +/- 0.3%; Cd: 1.4+/- 2.5%). Exposure of PT cells with Cd decreased protein levels of the catalytic subunit (alpha1) of Na+/K(+)-ATPase, a long-lived membrane protein (t(1/2)>48 h) that drives reabsorption of ions and nutrients through Na(+)-dependent transporters in PT. Incubation of PT cells for 48 h with Cd decreased Na+/K(+)-ATPase alpha1-subunit, as determined by immunoblotting, by approximately 50%, and NAC largely prevented this effect. Inhibitors of the proteasome such as MG-132 (20 microM) or lactacystin (10 microM), as well as lysosomotropic weak bases such as chloroquine (0.2 mM) or NH(4)Cl (30 mM), significantly reduced the decrease of Na(+)/K(+)-ATPase alpha1-subunit induced by Cd, and in combination abolished the effect of Cd on Na+/K(+)-ATPase. Immunofluorescence labeling of Na+/K(+)-ATPase showed a reduced expression of the protein in the plasma membrane of Cd-exposed cells. After addition of lactacystin and chloroquine to Cd-exposed PT cells, immunoreactive material accumulated into intracellular vesicles. The data indicate that micromolar concentrations of Cd can increase ROS production and exert a toxic effect on PT cells. Oxidative damage increases the degradation of Na+/K(+)-ATPase through both the proteasomal and endo-/lysosomal proteolytic pathways. Degradation of oxidatively damaged Na+/K(+)-ATPase may contribute to the 'Fanconi syndrome'-like Na(+)-dependent transport defects associated with Cd-nephrotoxicity.     

The beneficial role of potassium in Cd-induced stress alleviation and growth improvement in Gladiolus grandiflora L.

Heavy metal contaminated agricultural soils are one of the most important constraints for successful cultivation of crops. The current research was conducted to evaluate the role of potassium (K) on plant growth and amelioration of cadmium (Cd) stress in Gladiolus grandiflora under greenhouse conditions. G. grandiflora corms were sown in media contaminated with 0 (C), 50 (Cd50) and 100 (Cd100) mg Cd kg^?1 soil. The plants growing in Cd-contaminated media exhibited reduced gas exchange attributes, chlorophyll (Chl) contents, vegetative and reproductive growth as compared to control. The plants raised in Cd contaminated media showed reduced nutrition yet higher Cd contents. However, supplementation of 60 mg Kg^?1 K in treated plants (C+K, Cd50+K and Cd100+K) improved quantity of total soluble protein and proline (Pro) along with activity of antioxidant enzymes including superoxide dismutase (SOD), catalase (CAT), peroxidase (POD) and ascorbate peroxidase (APX) under Cd stress. Similarly, K supplementation reduced the level of malondialdehyde (MDA) and hydrogen peroxide (H₂O₂) in treated plants. Potassium supplemented plants exhibited better vegetative and reproductive growth. The improved stress tolerance in K supplemented plants was attributed to the reduced quantity of MDA and H₂O₂, enhanced synthesis of protein, proline, phenols, flavonides and improved activity of antioxidant enzymes. The present research supports the application of K for alleviation of Cd stress in G. grandiflora.     

Heavy metals content and distribution in basil (Ocimum basilicum L.) as influenced by cadmium and different potassium sources

Abstract

Despite the fact that cadmium (Cd) is a non-essential element for plants, it can influence nutrients and affect human health. Potassium (K) can influence the transportation of heavy metals (HMs) in soil-plant systems. Here, a greenhouse experiment was conducted to evaluate the effect of Cd and K fertilizers on the different partitioning forms of HMs, their concentrations, uptake in the shoots and roots of Ocimum basilicum. Treatments comprised 2 levels of Cd (0 and 40?mg kg^?1) and three levels of K (0, 100, and 200?mg kg^?1) from three sources, i.e. KCl, K₂SO_4, and K-nano-chelate. 40?mg Cd kg^?1 increased the shoot (above ground parts) Cd concentration. Addition of K as KCl, K₂SO_4, and K-nano-chelate increased the presence of Cd in shoots by 86, 82 and 76%, respectively, compared to the control. Using the nano-chelate of K can increase the accumulation of Cd in plants grown on contaminated soils to lesser content than that of the other forms of K. Application of 40?mg Cd kg^?1 reduced the concentration of Zn, Cu, and Mn in the shoot, but increased shoot Fe concentration. Transfer factor (TF), which is the ratio of metal concentration in shoot to its concentration in root, of the studied HMs, was significantly affected by Cd and K treatments. Therefore, the proper form and dose of chemical fertilizers should be applied in Cd-contaminated soils.     

The decline in potassium concentration is associated with cadmium toxicity of rice seedlings

Cadmium (Cd) is one of the most dangerous environmental pollutants, among other things, affecting plant mineral composition. Thus, in this study, we investigated the changes in potassium (K) concentration in Cd-treated rice (Oryza sativa L.) seedlings of two cultivars. On treatment with 5 μM CdCl₂, the Cd concentration increased in the shoot and roots of Cd-sensitive cultivar (cv. Taichung Native 1, TN1) but not or slightly in the Cd-tolerant cultivar (cv. Tainung 67, TNG67). The decrease in K concentration in the shoot and roots of TN1 caused by Cd was more pronounced than that of TNG67. Exogenous addition of KCl decreased Cd concentration and reduced Cd toxicity of TN1 seedlings. Evidence presented in this study suggests that the improvement of K status is able to reduce toxicity of rice seedlings to CdCl₂.