外源硅对甘蓝镉毒害的缓解机制研究

通过水培实验,以耐镉(Cd)性不同的甘蓝品种‘绿丰’(Cd敏感性)和‘牛心’(Cd耐性)为材料,研究外源硅(Si)对Cd胁迫下甘蓝生长、Cd含量、养分吸收、光合特性和抗氧化酶活性的影响,以揭示外源硅对甘蓝镉毒害的缓解作用机制。结果表明:(1)与对照相比,Cd胁迫显著抑制了甘蓝植株的生长,降低了植株体内营养元素(Mg、Fe、Mn、Cu)的含量以及抗氧化酶[超氧化物歧化酶(SOD)、抗坏血酸过氧化物酶(APX)、过氧化氢酶(CAT)]的活性和叶片的光合效率;并且显著增加了甘蓝植株内丙二醛(MDA)的含量。(2)与单独Cd处理相比,外源Si显著提高了甘蓝植株的生物量、地上部营养元素的含量以及光合参数[净光合速率(P_n)、气孔导度(G_s)及蒸腾速率(Tr)]、荧光参数值[原初光能转化效率(F_v/F_m)、实际光化学效率(Φ_(PSⅡ))、光化学淬灭系数(qP)]、光合色素含量和抗氧化酶活性;同时显著降低了甘蓝植株内MDA含量及Cd由根系向地上部的转运。研究发现,在镉胁迫环境下,外源Si可以通过降低Cd在甘蓝地上部的含量,提高抗氧化酶活性来降低膜脂过氧化,促进养分的吸收和转运,增强光合作用效率,从而缓解Cd对甘蓝生长的毒害。     

黄瓜中硅的生理功能及转运机制研究进展

硅是植物体的重要组成部分,尽管硅尚未被列为植物生长的必需元素,但它在促进植物生长发育、提高作物对非生物逆境（干旱、盐分和重金属等）和生物逆境（病虫害）抗性等方面都具有重要作用。硅不仅能改善植株对矿质营养的吸收,提高作物产量和品质,而且能沉积在叶片及叶鞘表皮细胞,形成硅化细胞和角质双硅层结构,增强寄主植物细胞壁的机械强度和稳固性,从而增强植物对真菌侵入和扩展的抵御能力,提高植物对金属离子毒害的抗性、缓解盐胁迫、增强抗高低温和抗紫外线辐射等。本文在植物硅素营养和转运机制研究的基础上,对硅素营养在黄瓜中生长发育、抗逆和吸收转运机制等方面的效应做了相关综述,并展望了黄瓜中硅研究的未来发展。     

外源水杨酸对镉胁迫下两种菊芋品系幼苗的缓解作用

研究了不同浓度Cd(0、50、100 μmol·L-1)及水杨酸(SA)(0、10、100和1000μmol·L-1)处理下2种菊芋(南芋5号和南芋2号)的生物量变化情况,测定了叶片色素(叶绿素和类胡萝卜素)含量及光合作用参数,并比较了2种菊芋对Cd的吸收转运差异性.结果表明:外源SA处理不同程度地缓解了Cd对菊芋幼苗生长的毒害效应,提高了色素含量,改善了光合作用参数,增大了Cd的生物富集系数(BCF)和转运系数(TF),与此同时减少了植株不同器官对Cd的累积量,但最佳SA浓度因品种及器官不同而有所区别.研究表明,适当浓度的SA处理可以有效增强2种菊芋对Cd的耐性.     

外源NO对镉胁迫下水稻幼苗抗氧化系统和微量元素积累的影响

重金属污染已成为世界范围的主要问题之一,其中镉(Cd)毒害的范围最广.Cd污染影响植物的生长和发育.一氧化氮(NO)作为植物体内的一种活性信号因子,参与了植物对各种胁迫的应答.为了探讨外源NO对Cd胁迫下水稻苗期生理生化响应的调节作用,以粳稻和籼稻为材料,采用营养液水培的方法,研究外源NO供体硝普钠(SNP)对100μmol/L Cd胁迫下2个水稻基因型(ZF802和ZH11)幼苗生长、抗氧化酶系统和微量元素吸收的影响.结果表明,1.5 mmol/L SNP能明显缓解Cd胁迫对两种水稻幼苗生长的抑制作用,提高幼苗对Cd的耐性.尽管存在一定的基因型差异,但外源NO能使两个基因型叶片中的超氧化物歧化酶(SOD)和过氧化物酶(POD)活性下降,降低了叶片中丙二醛(MDA)含量,从而缓解Cd对水稻幼苗的毒害.另外,外源NO影响Cd胁迫下水稻幼苗地上部和根部Cd和微量元素的积累,这种影响的程度与水稻的品种有关,具有显著的基因型差异,且机制较复杂.     

植物对重金属镉的耐受机制

镉离子（Cd^2＋）具有强植物毒性,抑制植物生长,甚至使植物死亡。由于长期的环境选择和适应进化,植物发展出耐受机制,可减轻或避免Cd^2＋的毒害。硫转运蛋白、硫还原相关酶类以及半胱氨酸、谷胱甘肽和植物螯合肽合成基因的表达受Cd^2＋调控。同时这些基因的过表达也能提高植物对Cd^2＋的耐性。植物抗氧化系统对Cd^2＋胁迫诱发的活性氧的清除作用,具转运Cd^2＋活性的质膜转运蛋白促进Cd^2＋经共质体途径向木质部运输、装载,而后随蒸腾流向地上部迁移,具转运Cd^2＋活性的液泡膜转运蛋白促进Cd^2＋进入液泡的隔离作用,都在植物对Cd^2＋的耐性中起作用。     

硒和碲对缓解糜子镉毒害及减少籽粒镉积累的调控效应

提高糜子(Panicum miliaceum)镉耐受性与低积累能力对镉污染地区糜子的安全生产具有重要意义。该研究以镉耐受和镉敏感糜子品种为材料, 通过苗期水培和全生育期盆栽试验, 分析不同形态硒和四价碲对镉胁迫下糜子生长、根系形态、镉吸收转运和籽粒矿质营养含量的影响。结果表明, 外源添加硒和碲能缓解镉毒害, 其中有机硒缓解效果较好。与单独镉处理相比, 硒和碲能够促进根系直径增加并抑制镉吸收, 最高可使根系镉含量降低33%。此外, 硒和碲能够增加细胞壁和液泡中镉的占比, 提高镉耐受性。叶面喷施硒提高了糜子籽粒中锌、锰和钼等矿质营养元素含量。无机四价硒能更有效地抑制镉从营养器官向籽粒的转运, 在5 mg·kg^-1镉处理下, 可使镉敏感和镉耐受品种籽粒镉含量分别降低11.3%和20.3%。综上, 外源添加硒能够显著提高糜子镉耐受性并减少籽粒镉积累。研究结果可为镉污染地区糜子的安全生产提供参考。     

铜、镉胁迫下外源NO介导的番茄解毒途径

一氧化氮(NO)作为信号分子,在抵御重金属胁迫中起重要作用,但对不同离子胁迫下的解毒机制尚缺乏研究.本研究采用营养液培养法,研究了铜(Cu)、镉(Cd)单一或复合胁迫下,番茄幼苗对Cu、Cd的吸收转运特性及对外源NO的响应机制.结果表明: 50 μmol·L^-1的Cu²⁺、Cd²⁺均显著抑制番茄植株的生长,其中Cd胁迫对生长的抑制效应远高于Cu胁迫.Cu、Cd单一或复合胁迫均使番茄根系Cu、Cd含量显著升高,但根系对Cu、Cd吸收存在严格选择性.根细胞对必需元素Cu表现出“奢侈吸收”的现象,而对毒性较强的Cd则吸收相对较少,胞内Cd浓度仅为Cu的1/10左右.外源NO处理可不同程度地缓解Cu、Cd胁迫,其中缓解Cd胁迫的效能更强.番茄对被动进入细胞的Cu、Cd具有相似的解毒机制:一方面,Cu、Cd胁迫诱导细胞质中产生谷胱甘肽(GSH)、植物螯合肽(PCs)和金属硫蛋白(MTs),络合过多的Cu、Cd离子,降低其生物毒性;另一方面,过多的Cu、Cd离子或螯合物被转运至液泡区隔化.外源NO通过调控GSH-GSSG(氧化型谷胱甘肽)氧化还原状态及GSH-PCs代谢方向的改变,促进Cu、Cd离子转运至液泡区隔化来缓解胁迫抑制;NO还可诱导植株叶片或根系表达更多的金属硫蛋白、GSH和PCs,而且上述响应普遍存在叠加效应.这可能是NO介导番茄对Cu、Cd胁迫的另一主要解毒途径.     

镉胁迫对不同品种小麦幼苗生长和生理特性的影响

以4个小麦品种(豫麦4、扬麦15、宝丰7228和宁麦13)为材料,研究了镉胁迫对水培条件下小麦幼苗生长和抗氧化酶系统的影响.结果表明:镉胁迫条件下植株的生长、叶绿素合成及硝酸还原酶活性明显受到抑制;植株体内与氧化胁迫有关的CAT和POD活性下降,H2O2和MDA含量上升;4个品种在镉处理后生长抑制和氧化胁迫程度上表现不同,显示出对镉耐性的差异,其中扬麦15对镉耐性高于其他3个品种.     

植物的硫同化及其相关酶活性在镉胁迫下的调节

植物对土壤中硫的利用包括根系对硫酸盐的吸收、转运、同化、分配等过程,也是由一系列酶和蛋白质参与和调节的代谢过程。近年来的研究表明,在植物体内,硫同化与植物对镉等重金属元素的胁迫反应机制有着密切关系。镉胁迫能调节植物对硫酸盐的吸收、转运、同化,以及半胱氨酸、谷胱甘肽（glutathione,GSH）和植物螯合肽（Dhytochelatins,pc）的合成。植物在镉胁迫下通过多种调节机制,增强对硫酸盐的吸收和还原,迅速合成半胱氨酸和谷胱甘肽等代谢物,从而合成足够的PC,以满足植物生理的需要。     

外源钙(Ca)对毛葱耐镉(Cd)胁迫能力的影响

以对环境敏感的毛葱(Allium cepa var.agrogarum L.)为材料,通过水培试验分别研究不同浓度Ca(0、0.1、1、10 mmol/L)对Cd(10μmol/L、100μmol/L和300μmol/L)胁迫下毛葱幼苗生长、叶片光合特性、体内Cd积累和矿质营养的变化,探讨Ca缓解敏感植物Cd毒害的生理生化机制。结果表明:(1)Cd显著抑制了毛葱的生长并导致其根端弯曲、发黄,叶片绿色加深;外源Ca显著削弱了Cd的毒害,缓解了其对毛葱生长的抑制;(2)Cd导致毛葱叶片光合色素含量大幅度上升却显著降低了毛葱的光合作用;外源Ca延缓了光合色素上升的速度,提高了毛葱的净光合速率、气孔导度和蒸腾速率;(3)Cd胁迫导致毛葱体内Cd含量显著增加,并造成明显的矿质营养失衡,主要表现为显著降低了毛葱根中Mg、Mn,叶中Ca、Mg、Mn、Zn等元素的含量,毛葱根中Ca、Fe、Zn,叶中Fe元素含量的显著增加,扰乱了毛葱体内矿质营养的内稳态;外源Ca削弱了毛葱对Cd的积累,减轻了Cd胁迫所造成的矿质营养失衡。因此,外源Ca能通过抑制Cd的吸收,促进叶片光合作用及气体交换速率,维持植物体的含水量、植物叶片光合色素含量及矿质营养的平衡等途径来增强毛葱对Cd胁迫的耐性。     

Cadmium absorption and transportation pathways in plants

Controlling the uptake, transport, translocation, and accumulation of excessive amounts of cadmium from polluted environments is critical for plants and, consequently, humans with regard to food safety. Plants adopt various cellular and molecular mechanisms to minimize Cd toxicity. Upon exposure to Cd, plants initially implement avoidance strategies, such as production of organic acids, chelation, and sequestration, to prevent metal access to root cells. Nevertheless, Cd can be transported through the roots, stems, and leaves via apoplastic and symplastic pathways. These processes have been controlled by specific sites at the root surface and root cortex, in cells responsible for loading the root xylem, at the transition between the vascular systems of the root and the shoot, and in connecting tissues and cells at the stem. Although resistance to heavy metal cadmium can be achieved by either avoidance or tolerance, genetic basis to tolerance is therefore implied, in that these mechanisms are heritable attributes of tolerant mutants or genotypes.     

重金属镉(Cd)在植物体内的转运途径及其调控机制

重金属镉(Cd)的毒害效应与其由土壤向植物地上部分运输有关,揭示Cd~(2+)转运途径及其调控机制可为提高植物抗镉性以及镉污染的植物修复提供依据。对Cd~(2+)在植物体内的转运途径,特别是限制Cd~(2+)移动的细胞结构和分子调控机制研究进展进行了回顾。Cd~(2+)通过共质体和质外体途径穿过根部皮层进入木质部的过程中,大部分在皮层细胞间沉积,少部分抵达中柱后转移到地上部分。为了免受Cd~(2+)的危害,植物体产生了多种限制Cd~(2+)吸收和转移的生理生化机制:1)环绕在内皮层径向壁和横向壁上的凯氏带阻止Cd~(2+)以质外体途径进入木质部;2)螯合剂与进入根的Cd~(2+)螯合形成稳定化合物并区隔在液泡中;3)通过H+/Cd~(2+)离子通道等将Cd~(2+)逆向转运出根部。植物共质体和质外体途径转运重金属镉的能力以及两条途径的串扰尚待进一步明晰和阐明。     

Exogenous nitric oxide improves antioxidative capacity and reduces auxin degradation in roots of Medicago truncatula seedlings under cadmium stress

The effects of nitric oxide (NO) on cadmium toxicity in Medicago truncatula seedlings were studied by investigating root growth and uptake of antioxidants, IAA and ions. Exposure to cadmium reduced root growth and NO accumulation, and increased the production of reactive oxygen species (ROS) in roots. Supplementation with NO improved root growth and reduced ROS accumulation in roots. The NO-scavenger cPTIO, the nitrate reductase (NR) inhibitor tungstate, and the NO synthase (NOS) inhibitor L-NAME all inhibited the accumulation of NO in roots and reversed the effects of NO in promoting the root growth and accumulation of proline and glutathione. Application of NO reduced auxin degradation by inhibiting the activity of IAA oxidase. Exogenous NO also enhanced the uptake of K⁺ and Ca²⁺. These results suggest that NO improves cadmium tolerance in plants by reducing oxidative damage, maintaining the auxin equilibrium and enhancing ion absorption.     

镉污染条件下水稻对假单胞菌TCd-1微生物修复的生理响应

微生物在修复重金属污染土壤中发挥重要作用。为阐明假单胞菌TCd-1降低水稻镉吸收的机理,通过加镉土培盆栽试验,在10 mg/kg镉处理下,对比研究了接种菌株对2种镉耐性不同水稻品种(特优671、百香139)镉含量、根系活力、光合作用、抗氧化酶活性及抗氧化物质含量的影响。结果表明：与10 mg/kg镉处理相比,接种假单胞菌TCd-1后水稻“特优671”和“百香139”的根、茎、叶、糙米的镉含量分别依次降低了30.4%、39.1%、40.7%、29.2%和52.2%、51.7%、18.4%、38.8%;提高了2种水稻的根系活力、光合作用,促进了水稻的生长;“特优671”、“百香139”叶片的超氧化物歧化酶(Superoxide dismutase, SOD)活性分别提高了7.3%、138.5%,过氧化物酶(Peroxidase, POD)活性提高了82.0%、106.2%,过氧化氢酶(Catalase, CAT)活性提高了58.8%、172.7%;叶片的类黄酮含量提高了139.4%、73.6%,总酚含量提高了27.2%、23.1%;超氧阴离子含量降低了44.2%、29.0%,丙二醛(Mal...     

Cadmium toxicity in crop plants and its alleviation by arbuscular mycorrhizal (AM) fungi: An overview

Cadmium (Cd), a toxic metal released into agricultural settings induces numerous changes in plant growth and physiology. The main known mechanisms of Cd toxicity include its affinity for sulfhydryl groups in proteins and its ability to replace some essential metals in active sites of enzymes, thus causing inhibition of enzyme activities and protein denaturation. This article reviews detrimental effects of Cd toxicity on the functional biology of plants and summarizes the mechanisms that are activated by plants to prevent the absorption or to detoxify Cd ions such as synthesis of antioxidants, osmolytes, phytochelatins, metallothioneins, etc. Arbuscular mycorrhizal (AM) fungi are reported to be present on the roots of plants growing in metal-contaminated soils and play an important role in metal tolerance. Through mycorrhizal symbiosis, heavy metals are immobilized in the rhizosphere through precipitation in the soil matrix, adsorption onto the root surface or accumulation within roots, and compartmentalized in aboveground parts of the plant. This article unfolds the potential role of AM fungi in enhancing Cd tolerance of plants.     

Role of salicylic acid in resistance to cadmium stress in plants

Key message

We review and introduce the importance of salicylic acid in plants under cadmium stress, and provide insights into potential regulatory mechanisms for alleviating cadmium toxicity.

Abstract

Cadmium (Cd) is a widespread and potentially toxic environmental pollutant, originating mainly from rapid industrial processes, the application of fertilizers, manures and sewage sludge, and urban activities. It is easily taken up by plants, resulting in obvious toxicity symptoms, including growth retardation, leaf chlorosis, leaf and root necrosis, altered structures and ultrastructures, inhibition of photosynthesis, and cell death. Therefore, alleviating Cd toxicity in plants is a major aim of plant research. Salicylic acid (SA) is a ubiquitous plant phenolic compound that has been used in many plant species to alleviate Cd toxicity by regulating plant growth, reducing Cd uptake and distribution in plants, protecting membrane integrity and stability, scavenging reactive oxygen species and enhancing antioxidant defense system, improving photosynthetic capacity. Furthermore, SA functions as a signaling molecule involved in the expression of several important genes. Significant amounts of research have focused on understanding SA functions and signaling in plants under Cd stress, but several questions still remain unanswered. In this article, the influence of SA on Cd-induced stress in plants and the potential regulation mechanism for alleviating Cd toxicity are reviewed.

     

Responses of nitric oxide and hydrogen sulfide in regulating oxidative defence system in wheat plants grown under cadmium stress

We conducted a study to evaluate the interactive effect of NO and H₂S on the cadmium (Cd) tolerance of wheat. Cadmium stress considerably reduced total dry weight, chlorophyll a and b content and ratio of Fv/Fm by 36.7, 48.6, 26.7 and 19.5%, respectively, but significantly enhanced the levels of hydrogen peroxide (H₂O₂) and malondialdehyde (MDA), endogenous H₂S and NO, and the activities of antioxidant enzymes. Exogenously applied sodium nitroprusside (SNP) and sodium hydrosulfide (NaHS), donors of NO and H₂S, respectively, enhanced total plant dry matter by 47.8 and 39.1%, chlorophyll a by 92.3 and 61.5%, chlorophyll b content by 29.1 and 27.2%, Fv/Fm ratio by 19.7 and 15.2%, respectively, and the activities of antioxidant enzymes, but lowered oxidative stress and proline content in Cd-stressed wheat plants. NaHS and SNP also considerably limited both the uptake and translocation of Cd, thereby improving the levels of some key mineral nutrients in the plants. Enhanced levels of NO and H₂S induced by NaHS were reversed by hypotuarine application, but they were substantially reduced almost to 50% by cPTIO (a NO scavenger) application. Hypotuarine was not effective, but cPTIO was highly effective in reducing the levels of NO and H₂S produced by SNP in the roots of Cd-stressed plants. The results showed that interactive effect of NO and H₂S can considerably improve plant resistance against Cd toxicity by reducing oxidative stress and uptake of Cd in plants as well as by enhancing antioxidative defence system and uptake of some essential mineral nutrients.     

褪黑素对玉米幼苗根系发育和抗旱性的影响

褪黑素是一种在生物体内广泛存在的吲哚胺类化合物,参与植物的多种生理和生化过程。近年来研究认为褪黑素可以不同程度地增强植物的抗逆性,但对其作用机理仍知之甚少。通过两种褪黑素的施用方法,详细研究了褪黑素对于玉米根系发育和抗旱性的影响。首先,采用水培根灌褪黑素的方法对玉米幼苗的根系和生长状况进行分析,结果表明施加褪黑素显著提高多种玉米幼苗根系参数,包括根长、根表面积、根体积和侧根数目等。其次,采用盆栽浸种褪黑素的方法,对叶片相对含水量、光合作用、抗氧化酶活性、地上部分生物量等进行测定,结果表明在干旱胁迫条件下,褪黑素浸泡种子的处理方式能够提高植株的光合速率、气孔导度和蒸腾速率,增强抗氧化酶活性以及降低活性氧和丙二醛含量,证明褪黑素促进植物根系发育,减轻氧化损伤,缓解光合抑制,改善植物水分状况,从而提高植物抗旱性。     

硅提高植物抗旱性的生理机制研究进展

干旱作为限制作物产量和品质的主要非生物胁迫之一,对全球社会、经济和生态造成巨大损失。在全球气候变化背景下,提高植物抗旱性的重要性日益突显。硅能够提高植物的抗旱性：外源硅的施用可以影响气孔导度,改变蒸腾速率,改善植物水分状况;通过调节气孔动力学、合成光合色素,促进光化学反应,从而改善光合作用;此外硅可通过渗透调节以平衡植物对矿质元素的吸收,以及调节抗氧化防御系统,减轻植物在干旱胁迫中的氧化损伤。总结了硅对干旱胁迫下植物水分利用、光合作用、矿质元素吸收、抗氧化系统、植物激素代谢等方面的作用及相关生理机制。建议未来从复合逆境胁迫、低硅积累植物等方面进一步揭示硅提高植物抗旱性的作用机制,从而为农林生态系统合理利用硅素来提高生产效率提供科学依据和理论基础。     

Cellular Mechanisms in Higher Plants Governing Tolerance to Cadmium Toxicity

Cadmium (Cd) is an inorganic mineral in the earth's crust. Cadmium entry into the environment occurs through geogenic and anthropogenic sources. Industrial activities including mining, electroplating, iron and steel plants, and battery production employ Cd during their processes and often release Cd into the environment. When disseminated into soil, Cd can be detrimental to agro-ecosystems because it is relatively mobile and phytotoxic even at low concentrations. Cadmium's phytotoxicity is due to reductions in the rate of transpiration and photosynthesis and chlorophyll concentration resulting in retardation of plant growth, and an alteration in the nutrient concentration in roots and leaves. In response to Cd toxicity, plants have developed protective cellular mechanisms such as synthesis of phytochelatins and metallothioneins, metal compartmentalization in vacuoles, and the increased activity of antioxidant enzymes to neutralize Cd-induced toxicity. While these direct protective mechanisms can help alleviate Cd toxicity, other indirect mechanisms such as microelements (zinc, iron, manganese, and selenium) interfering with Cd uptake may decrease Cd concentration in plants. This comprehensive review encompasses the significance of Cd, portals of contamination and toxicity to plants, and implications for crop production. Various mitigation strategies with the beneficial effects of zinc, iron, manganese, and selenium in activating defence mechanisms against Cd stress are discussed. Furthermore, this review systematically identifies and summarises suitable strategies for mitigating Cd-induced toxicity in plants.