甘蓝（Brassica oleracea L.）耐镉性的品种差异及其机理研究
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摘要
随着工业和农业的迅速发展,土壤中的重金属污染已经成为当今世界上主要的环境问题之一。Cd易在动、植物器官中积累且不易被分解,被认为是对人类最具毒性的污染物质之一。在中国,叶菜类蔬菜是仅次于谷类的人们重要的日常食物,且其具有较强的积累重金属的能力。目前有关作物Cd耐性、吸收与积累品种间差异的机理尚不完全清楚,从而限制了相关育种与栽培工作的开展。本研究在明确甘蓝(Brassica oleracea L.) Cd耐性和Cd含量品种间差异的基础上,利用水培实验,系统地研究Cd抑制甘蓝生长和Cd耐性、吸收与积累品种间差异的生理机制,并探索减少甘蓝Cd胁迫和吸收的化学调控技术。
1、不同甘蓝品种对Cd胁迫的响应
采用盆栽试验,比较研究了31个甘蓝品种对Cd浓度的反应。结果表明,100mg·kg-1Cd处理下,不同甘蓝品种的耐性及镉吸收积累能力具有显著的品种间差异。地上部干重的抑制率变幅为-15.4%～91．3%,镉对地上部的抑制作用大于根。根系Cd含量变幅为617～1789mg·kg-1,地上部的Cd含量变幅为47～183mg·kg-1,根系Cd含量显著高于地上部。初步筛选出10个对Cd耐性不同的甘蓝品种进行50μmo·L-1Cd的溶液处理,根据植株对Cd的耐性和从根部向地上部转运Cd的能力,筛选出春丰(Cd-耐性品种、低转运Cd)和绿丰(Cd-敏感品种、高转运Cd)2个品种进行甘蓝耐Cd机理的研究。
2、镉胁迫对甘蓝生长和镉吸收的影响
为了研究甘蓝的耐Cd机理,以2个Cd耐性不同的甘蓝品种(春丰,耐Cd品种；绿丰,Cd敏感品种)为试材,比较研究了不同Cd浓度(0、20、50、100μmol·L-1对甘蓝植株生长、Cd吸收和植株内Cd分布的影响。结果表明,20μmol·L-1的Cd对甘蓝根系伸长及植株干重产生显著的抑制作用,但春丰具有更强的耐Cd胁迫的能力。Cd主要分布在甘蓝的根系。春丰叶中的Cd含量显著低于绿丰,而根中Cd含量显著高于绿丰。绿丰和春丰吸收的Cd约70%-74%和66%-68%被转运到地上部。2个品种叶、茎和根系中的Cd大部分结合在细胞壁上,且春丰细胞壁上的Cd含量显著高于绿丰。与绿丰相比,春丰对Cd的耐性较强可能归因于春丰有更强的阻遏Cd向地上部运输和将Cd结合在细胞壁上的能力。
3、镉胁迫对甘蓝光合特性和养分吸收的影响
Cd敏感品种绿丰在低浓度Cd (20μmol·L-1)处理下表现出明显受抑,叶片净光合速率(Pn)、气孔导度(Gs)、蒸腾速率(Ti)、PSⅡ光化学效率(Fv/Fm)及PS II光合电子传递的量子效率(ΦDPS Ⅱ)显著降低；而Cd耐性品种春丰在高浓度Cd(50μmol·L-1和100μmol·L-1)处理下才受到显著影响,且相同的Cd水平下,春丰受抑程度显著低于绿丰。Cd胁迫降低了2品种叶绿素a和b含量,尤以对叶绿素a的的影响更甚,进而抑制了叶片光合能力。Cd胁迫显著抑制了2品种对Mn的吸收,促进了对Zn和S的吸收,抑制了Zn和S(绿丰)从根系向地上部的转运,且绿丰受抑幅度更大。此外,Cd处理还显著抑制了绿丰对Fe、Mg的吸收,降低了其叶片中Fe、Mg的含量,而对春丰无显著影响。Cd促进了春丰对P的吸收,而绿丰正相反。因此,与耐性品种春丰相比,Cd胁迫下敏感品种绿丰叶片中Mn、Fe、 Mg、Zn、Ca和P的含量显著降低,是影响其叶片光合作用,进而抑制植株生长的重要生理原因。
4、镉胁迫对甘蓝膜脂过氧化及抗氧化酶系统的影响
过量Cd会诱导植物体内活性氧(ROS)的产生,但植物体内的抗氧化防卫系统能够在一定范围内清除ROS。耐性品种春丰和敏感品种绿丰分别在Cd处理后第4d和2d,植株内的MDA含量没有受到显著影响,表明抗氧化酶系统在清除ROS中起了重要作用,其中主要是SOD和POD活性显著升高。除春丰叶片中GR活性持续增强外,2品种叶片和根系中SOD、POD、CAT、APX和GR活性随Cd处理时间的延长均呈先升后降的趋势,而MDA含量逐步增加,表明ROS的产生已经超过了防御系统的清除能力,产生了膜脂过氧化。与春丰相比,绿丰体内抗氧化酶活性短时间内急剧升高而又显著下降,至Cd处理第10d,各酶活性均显著低于春丰,而MDA含量显著高于春丰,说明绿丰受到了更大的氧化胁迫。可以认为,Cd处理诱导了ROS的产生,造成了膜脂过氧化,这是Cd伤害甘蓝植株的重要机制之一。抗氧化能力的不同,能部分解释春丰和绿丰对Cd的耐性差异。
5、非蛋白巯基和有机酸在甘蓝耐镉性中的作用
与对照相比,Cd处理显著增加了2个甘蓝品种叶片和根系中非蛋白巯基(NPT)、植物螯合肽(PCs)、柠檬酸和脯氨酸的含量,且耐Cd品种春丰增加的幅度显著高于敏感品种绿丰。与绿丰相比,春丰对Cd的耐性较强也许归因于其金属配位体,如PCs和柠檬酸的含量较高及渗透调节能力较强。然而,PCs和柠檬酸对解Cd毒的贡献也许小于细胞壁对Cd的固定(结合第三章的结果)。
6、根系细胞壁在甘蓝耐镉性中的作用
先前的研究已明确,根系细胞壁在甘蓝抗Cd胁迫中起重要作用。为了进一步研究甘蓝耐Cd机理,我们对2品种根系细胞壁成分进行了分析。结果表明：Cd胁迫显著增加了敏感品种绿丰细胞壁中果胶、半纤维素I和半纤维素II的含量及耐Cd品种春丰细胞壁中果胶和半纤维素II的含量。与绿丰相比,春丰细胞壁多糖含量增幅较小。Cd处理下,春丰果胶、半纤维素I和半纤维素II中糖醛酸的含量及绿丰果胶和半纤维素II中糖醛酸的含量均显著增加,而绿丰半纤维素I中糖醛酸的含量没有明显的变化。春丰糖醛酸含量显著高于绿丰,说明糖醛酸在甘蓝耐Cd性中起重要作用。2品种细胞壁各成分中的Cd含量以果胶中最高,它可能是Cd‘的主要结合位点。耐性品种春丰果胶中的Cd含量显著高于敏感品种绿丰,其糖醛酸残基与Cd的结合,可能是春丰耐Cd毒害的一个重要机制。
7、外源一氧化氮供体硝普钠(SNP)对甘蓝镉毒害的缓解效应
以Cd-敏感品种(绿丰)和Cd-耐性品种(春丰)为试料,研究了100μmol·L-1硝普纳(SNP, NO供体)对种子萌发、幼苗生长和生理的影响。结果表明：20-400μmol·L-1Cd单独处理抑制了2品种种子的萌发,50μmol·L-1Cd降低了植株的干物质重和叶绿素含量,绿丰受Cd的毒害更为明显。外源SNP同时处理下,Cd对甘蓝生长的抑制得到恢复。SNP对Cd胁迫下甘蓝植株的保护作用表现在：(a)显著降低了植株对Cd的吸收,明显减少Cd在叶片和根系细胞器中的积累；(b)降低了被Cd诱导增加的根系细胞壁多糖和多糖中糖醛酸的含量；(c)增强抗氧化酶活性和抗氧化物质的含量,降低了丙二醛(MDA)和过氧化氢(H202)的含量,保护了膜系统的完整性；(d)Cd诱导的非蛋白巯基(NP-SH)的增加被进一步提高,然而PCs的含量略有下降,表明GSH在保护2品种植株抵抗Cd毒害中起重要作用；(e)促进了根系对营养物质的吸收,增加了营养元素在叶片中的分布。Cd诱导S含量的增加被进一步增强,推测S的增加可能与NP-SH的合成有关；(f) SNP同时处理减轻了Cd对甘蓝净光合速率(Pn)的抑制,可能部分缘于SNP对光合机构的保护(提高了叶绿素含量、Fv/Fm值和ΦPSⅡ,增强了PSI I的活性)。综上所述,SNP对Cd胁迫下甘蓝植株的保护可能源于植物体内复杂的信号转导,交叉调控的生理和分子机制。
The contamination of soils with heavy metals is becoming a major environmental concern with the rapid development of agriculture and industries throughout the world. Cadmium (Cd) is considered to be one of the most toxic pollutants for humans due to its longevity and accumulation in their organs. Due to its high mobility in the soil-plant system, Cd is readily taken up by the roots and transported to other parts of plants. Leafy vegetables are an important component of human diet after cereals in China. They have an ability to take up heavy metals from contaminated soils and transport these metals to the shoots, which may lead to a risk for the health of the people. However, there is little information about varietal difference in Cd tolerance, uptake and accumulation, and which has in some extent limited the developments of its related breeding and cultivation programs to prevent Cd accumulation in crops. The investigation was carried out to study the physiological mechanism of varietal difference in Cd-uptake, accumulation and tolerance of cabbage and detrimental effects of Cd on growth, based on the research of varietal variation of Cd tolerance and Cd concentration, by using four levels of Cd hydroponic experiments. Meanwhile, the possibility of reducing Cd uptake and accumulation in cabbage by application of chemical regulators was also studied in the investigation. The major results were summarized as follows:
1. Response of different varieties of cabbage to cadmium stress
Pot experiment was conduced to investigate the reaction of31cabbage varieties to Cd toxicity. The results showed that these varieties had significant difference in Cd tolerance, uptake and accumulation when plants grew in the soil contaminated100mg·kg-1Cd. Among the varieties, the inhibition percentage of shoots dry weight varied from-15.4%to91.3%. The detrimental effects of Cd on shoots growth exceeded on roots. The range of Cd concentration in roots was from617to1789mg·kg-1and in shoots from47to183mg·kg-1, which roots exceeded shoots. Based on pot experiment,10varieties screened preliminarily as Cd-tolerance and Cd-sensitivity respectively were planted in a solution with50μmo·L-1 Cd. According to plant tolerance to Cd and the capacity to transport Cd from the roots to the shoots, Chunfeng and lufeng were eventually chosen as the Cd resistant and low Cd transportion variety and the Cd sensitive and high Cd transportion variety, respectively, to investigate the Cd tolerance mechanisms in cabbage.
2. Effects of Cd on growth and Cd uptake of cabbage seedlings
In order to study possible Cd resistance mechanisms in cabbage, several parameters on metal uptake and distribution were compared between the variety Chunfeng (Cd-resistance) and Lufeng (Cd-sensitive).The results showed that low Cd (20μM) exposure for both varieties caused a sharp decline (P<0.05) in the root elongation and the dry weight of plant, with a much more severe responsein the Cd-sensitive varieties. Cd taken up by cabbage plant was mainly distributed in roots compared to shoots, and the variety Chunfeng contained significantly lower Cd concentrations in the leaves and higher Cd ones in roots than Lvfeng. About70-74%and66-68%of the Cd taken up by Lvfeng and Chunfeng, respectively, was transported to shoots. The majority of the Cd accumulated in the leaves, stems and roots of both varieties appeared to be bound on the cell walls. The higher Cd resistance in Chunfeng than in Lufeng may be attributed to higher capacity to limit Cd uptake into shoots and to complex Cd on cell walls.
3. Effects of Cd on photosynthesis, chlorophyll fluorescence and nutrient uptake of cabbage seedlings
Cadmium stress impacted photosynthesis and chlorophyll fluorescence of cabbage plants. Low Cd (20μM) exposure for the Cd-sensitive variety Lufeng caused a sharp decline (P<0.05) in the net photosynthetic rate (Pn), stomatal conductance (Gs), photochemical efficiency of PS Ⅱ (Fv/Fm) and quantum yield of electron flow through PS Ⅱ (OPS Ⅱ) compared to the control. However, these deteriorative effects for the Cd-tolerant variety Chunfeng could be found only at high Cd (50～100μM) levels. At same Cd level, the inhibitions of these parameters were more pronounced in Lufeng than in Chunfeng. Cd stress reduced the concentration of chlorophyll a and chlorophyll b, especially for chlorophyll b, which could be an important factor of photosynthesis inhibition. For these two varieties, Cd stress significantly inhibited the Mn uptake, promoted Zn and S uptake, restrained the transportion of Zn and S (for Lufeng) from roots to shoots, and the decreases of nutrient uptake in lufeng were more obvious than in Chunfeng. Moreover, Cd stress also significantly inhibited Liifeng on Fe and Mg uptake, reduced its Fe and Mg concentration in leaves, but not significant effect on Chunfeng. Cd stress promoted the P uptake by Chunfeng, but opposite results were observed in Lufeng. Therefore, physiologically, the decrease of Mn, Fe, Mg, Zn、Ca and P concentrations in leaves of Lufeng under Cd stress was the key reason for the restraint of leaf photosynthesis, and the decrease of plant growth.
4. Effects of Cd on lipid peroxidation and activities of antioxidant capacity of cabbage seedlings
Excess Cd can cause an increased formation of reactive oxygen species (ROS). However, plants have evolved antioxidantive defence systems to keep ROS under control. The concentration of MDA in Lufeng and Chunfeng plants had not been significantly affected2d and4d, respectively, after Cd treatments, suggesting the antioxidantive defence systems played an importance role, mainly SOD and POD activities significantly increased. Apart from GR activity in leaves of Chunfneg increasing, the SOD, POD, CAT, APX and GR in leaves and roots of two varieties all increased firstly and then decreased with extended Cd exposure of time, but the concentration of MDA significantly increased, which indicated that the increased formation of ROS had exceeded the capacity of antioxidantive defence systems to scavenge ROS, resulting in membrane lipid peroxidation. The activities of antioxidant enzymes in Lufeng plants rised sharply within a short time and then declined significantly compared to that in Chunfeng, to the10th day exposed to Cd, the enzymes all were significantly lower than that in Chunfeng, but the concentration of MDA markedly higher than in Chunfeng, indicating that Lufeng was subjected to a greater degree of oxidative stress. Taken together, Cd induces the formation of ROS, resulting in a lipid peroxidation, which was an important mechanism of Cd toxicity to cabbage plants. The difference of antioxidant capacity might partly explain the Cd resistance of Chunfeng and Lufeng.
5. Role of non-protein thiols and organic acid in Cd tolerance of cabbage
A hydroponic experiment was carried out to examine the role of non-protein thiols and organic acid in Cd tolerance of cabbage. The results showed that compared with the control groups, Cd treatment significantly increased concentrations of non-protein thiol, PCs, citric acids and proline in the leaves and roots of two varieties. The increases were more pronounced in Chunfeng than in Lufeng. Taken together, it is suggested the higher Cd resistance in Chunfeng than in Lvfeng may be attributed to metal binding ligands such as PCs and citric acid, and osmotic adjustment ability. However, the contributions of SH-PCs and citric acid to Cd detoxification might be smaller than that of the cell walls.
6. Role of root cell wall in Cd tolerance of cabbage
Previous research has showed that the cell wall of roots played a important role in cabbage resisting Cd toxicity. To further understand the Cd resistance mechanism in cabbage, changes in cell wall properties of roots under Cd treatments were examined. The results showed that Cd stress evidently increased the concentrations of pectin, hemicellulose I and hemicellulose II in the cell wall of Cd-sensitive variety Liifeng and pectin and hemicellulose II in that of Cd-resistant one Chunfeng. The increase was more pronounced in Lufeng than in Chunfeng. The concentration of the uronic acid in pectin, hemicellulose I and hemicellulose II in Chunfeng and in the pectin and hemicellulose II in Lufeng increased markedly under Cd treatments, but not vary much in hemicellulose I in Lufeng. Chunfeng contained significant higher the uronic acid content than Liifeng, which suggested an important role of the uronic acid in Cd tolerance in cabbage. In addition, among the cell wall polysaccharides in two varieties roots, the Cd content in pectin was the highest, suggesting the majority of the Cd accumulated in root cell walls was tightly bound to the pectin. The Cd concentration in pectin of the variety Chunfeng was significantly higher than that of the variety Lufeng. The results suggested the higher Cd resistance in Chunfeng than in Lvfeng may be attributed to Cd-bound on the uronic acid in pectin.
7. Sodium nitroprusside (as nitric oxide donator) supplementation ameliorates cadmium-toxicity in cabbage
A hydroponic experiment was performed using Cd-sensitive (var. Lufeng) and Cd-tolerant (var. Chunfeng) cabbage seedling to evaluate how different varieties responsed to Cd toxicity in the presence of sodium nitroprusside (SNP), a nitric oxide (NO) donor. Results showed that20-400μM Cd levels inhibited seed germination, and50μM Cd reduced the biomass and chlorophyll concent, with a much more severe response in the Cd-sensitive variety. Cd+SNP treatment restored the Cd-inhibited growth. Exogenous SNP dramatically alleviated Cd toxicity, mainly observed:(a) markedly reduced Cd uptake by plants, obviously diminished Cd accumulation in subcellular fraction of leaves and roots of two varieties,(b) restored the increase of Cd-induced polysaccharides and uronic acid in polysaccharides contents in cell walls of roots,(c) enhanced the activities of antioxidant enzymes and the concentration of antioxidant, whereas decreased the concentrations of malondialdehyde (MDA) and hydrogen peroxide (H2O2), and helped to maintain stability and integrity of membrane system.(d) elevated the increased NP-SH contents, whereas declined slightly in the PCs content, which suggested the GSH may play a important role in protecting cabbage plants from Cd toxicity.(e) promoted nutrients uptake and increased nutrient elements distribution in leavese, and upregulated Cd-induced increase in S content in both varieties, inferring S increment may be involved in NP-SH synthesis;(f) Exogenous SNP reduced the inhibition of Cd on net photosynthetic rate (P n), which might partly be attributed to the NO protection to photosynthetic apparatus such as increasing the chlorophyll content, Fv/Fm, OPS Ⅱ and PSI Ⅰ activity. Taken together, these data suggest that the protection of NO to cabbage seedling against Cd toxicity may be involved in complex signal transduction in plants and cross regulation of physiological and molecular mechanisms.
1、不同甘蓝品种对Cd胁迫的响应
采用盆栽试验,比较研究了31个甘蓝品种对Cd浓度的反应。结果表明,100mg·kg-1Cd处理下,不同甘蓝品种的耐性及镉吸收积累能力具有显著的品种间差异。地上部干重的抑制率变幅为-15.4%～91．3%,镉对地上部的抑制作用大于根。根系Cd含量变幅为617～1789mg·kg-1,地上部的Cd含量变幅为47～183mg·kg-1,根系Cd含量显著高于地上部。初步筛选出10个对Cd耐性不同的甘蓝品种进行50μmo·L-1Cd的溶液处理,根据植株对Cd的耐性和从根部向地上部转运Cd的能力,筛选出春丰(Cd-耐性品种、低转运Cd)和绿丰(Cd-敏感品种、高转运Cd)2个品种进行甘蓝耐Cd机理的研究。
2、镉胁迫对甘蓝生长和镉吸收的影响
为了研究甘蓝的耐Cd机理,以2个Cd耐性不同的甘蓝品种(春丰,耐Cd品种；绿丰,Cd敏感品种)为试材,比较研究了不同Cd浓度(0、20、50、100μmol·L-1对甘蓝植株生长、Cd吸收和植株内Cd分布的影响。结果表明,20μmol·L-1的Cd对甘蓝根系伸长及植株干重产生显著的抑制作用,但春丰具有更强的耐Cd胁迫的能力。Cd主要分布在甘蓝的根系。春丰叶中的Cd含量显著低于绿丰,而根中Cd含量显著高于绿丰。绿丰和春丰吸收的Cd约70%-74%和66%-68%被转运到地上部。2个品种叶、茎和根系中的Cd大部分结合在细胞壁上,且春丰细胞壁上的Cd含量显著高于绿丰。与绿丰相比,春丰对Cd的耐性较强可能归因于春丰有更强的阻遏Cd向地上部运输和将Cd结合在细胞壁上的能力。
3、镉胁迫对甘蓝光合特性和养分吸收的影响
Cd敏感品种绿丰在低浓度Cd (20μmol·L-1)处理下表现出明显受抑,叶片净光合速率(Pn)、气孔导度(Gs)、蒸腾速率(Ti)、PSⅡ光化学效率(Fv/Fm)及PS II光合电子传递的量子效率(ΦDPS Ⅱ)显著降低；而Cd耐性品种春丰在高浓度Cd(50μmol·L-1和100μmol·L-1)处理下才受到显著影响,且相同的Cd水平下,春丰受抑程度显著低于绿丰。Cd胁迫降低了2品种叶绿素a和b含量,尤以对叶绿素a的的影响更甚,进而抑制了叶片光合能力。Cd胁迫显著抑制了2品种对Mn的吸收,促进了对Zn和S的吸收,抑制了Zn和S(绿丰)从根系向地上部的转运,且绿丰受抑幅度更大。此外,Cd处理还显著抑制了绿丰对Fe、Mg的吸收,降低了其叶片中Fe、Mg的含量,而对春丰无显著影响。Cd促进了春丰对P的吸收,而绿丰正相反。因此,与耐性品种春丰相比,Cd胁迫下敏感品种绿丰叶片中Mn、Fe、 Mg、Zn、Ca和P的含量显著降低,是影响其叶片光合作用,进而抑制植株生长的重要生理原因。
4、镉胁迫对甘蓝膜脂过氧化及抗氧化酶系统的影响
过量Cd会诱导植物体内活性氧(ROS)的产生,但植物体内的抗氧化防卫系统能够在一定范围内清除ROS。耐性品种春丰和敏感品种绿丰分别在Cd处理后第4d和2d,植株内的MDA含量没有受到显著影响,表明抗氧化酶系统在清除ROS中起了重要作用,其中主要是SOD和POD活性显著升高。除春丰叶片中GR活性持续增强外,2品种叶片和根系中SOD、POD、CAT、APX和GR活性随Cd处理时间的延长均呈先升后降的趋势,而MDA含量逐步增加,表明ROS的产生已经超过了防御系统的清除能力,产生了膜脂过氧化。与春丰相比,绿丰体内抗氧化酶活性短时间内急剧升高而又显著下降,至Cd处理第10d,各酶活性均显著低于春丰,而MDA含量显著高于春丰,说明绿丰受到了更大的氧化胁迫。可以认为,Cd处理诱导了ROS的产生,造成了膜脂过氧化,这是Cd伤害甘蓝植株的重要机制之一。抗氧化能力的不同,能部分解释春丰和绿丰对Cd的耐性差异。
5、非蛋白巯基和有机酸在甘蓝耐镉性中的作用
与对照相比,Cd处理显著增加了2个甘蓝品种叶片和根系中非蛋白巯基(NPT)、植物螯合肽(PCs)、柠檬酸和脯氨酸的含量,且耐Cd品种春丰增加的幅度显著高于敏感品种绿丰。与绿丰相比,春丰对Cd的耐性较强也许归因于其金属配位体,如PCs和柠檬酸的含量较高及渗透调节能力较强。然而,PCs和柠檬酸对解Cd毒的贡献也许小于细胞壁对Cd的固定(结合第三章的结果)。
6、根系细胞壁在甘蓝耐镉性中的作用
先前的研究已明确,根系细胞壁在甘蓝抗Cd胁迫中起重要作用。为了进一步研究甘蓝耐Cd机理,我们对2品种根系细胞壁成分进行了分析。结果表明：Cd胁迫显著增加了敏感品种绿丰细胞壁中果胶、半纤维素I和半纤维素II的含量及耐Cd品种春丰细胞壁中果胶和半纤维素II的含量。与绿丰相比,春丰细胞壁多糖含量增幅较小。Cd处理下,春丰果胶、半纤维素I和半纤维素II中糖醛酸的含量及绿丰果胶和半纤维素II中糖醛酸的含量均显著增加,而绿丰半纤维素I中糖醛酸的含量没有明显的变化。春丰糖醛酸含量显著高于绿丰,说明糖醛酸在甘蓝耐Cd性中起重要作用。2品种细胞壁各成分中的Cd含量以果胶中最高,它可能是Cd‘的主要结合位点。耐性品种春丰果胶中的Cd含量显著高于敏感品种绿丰,其糖醛酸残基与Cd的结合,可能是春丰耐Cd毒害的一个重要机制。
7、外源一氧化氮供体硝普钠(SNP)对甘蓝镉毒害的缓解效应
以Cd-敏感品种(绿丰)和Cd-耐性品种(春丰)为试料,研究了100μmol·L-1硝普纳(SNP, NO供体)对种子萌发、幼苗生长和生理的影响。结果表明：20-400μmol·L-1Cd单独处理抑制了2品种种子的萌发,50μmol·L-1Cd降低了植株的干物质重和叶绿素含量,绿丰受Cd的毒害更为明显。外源SNP同时处理下,Cd对甘蓝生长的抑制得到恢复。SNP对Cd胁迫下甘蓝植株的保护作用表现在：(a)显著降低了植株对Cd的吸收,明显减少Cd在叶片和根系细胞器中的积累；(b)降低了被Cd诱导增加的根系细胞壁多糖和多糖中糖醛酸的含量；(c)增强抗氧化酶活性和抗氧化物质的含量,降低了丙二醛(MDA)和过氧化氢(H202)的含量,保护了膜系统的完整性；(d)Cd诱导的非蛋白巯基(NP-SH)的增加被进一步提高,然而PCs的含量略有下降,表明GSH在保护2品种植株抵抗Cd毒害中起重要作用；(e)促进了根系对营养物质的吸收,增加了营养元素在叶片中的分布。Cd诱导S含量的增加被进一步增强,推测S的增加可能与NP-SH的合成有关；(f) SNP同时处理减轻了Cd对甘蓝净光合速率(Pn)的抑制,可能部分缘于SNP对光合机构的保护(提高了叶绿素含量、Fv/Fm值和ΦPSⅡ,增强了PSI I的活性)。综上所述,SNP对Cd胁迫下甘蓝植株的保护可能源于植物体内复杂的信号转导,交叉调控的生理和分子机制。
The contamination of soils with heavy metals is becoming a major environmental concern with the rapid development of agriculture and industries throughout the world. Cadmium (Cd) is considered to be one of the most toxic pollutants for humans due to its longevity and accumulation in their organs. Due to its high mobility in the soil-plant system, Cd is readily taken up by the roots and transported to other parts of plants. Leafy vegetables are an important component of human diet after cereals in China. They have an ability to take up heavy metals from contaminated soils and transport these metals to the shoots, which may lead to a risk for the health of the people. However, there is little information about varietal difference in Cd tolerance, uptake and accumulation, and which has in some extent limited the developments of its related breeding and cultivation programs to prevent Cd accumulation in crops. The investigation was carried out to study the physiological mechanism of varietal difference in Cd-uptake, accumulation and tolerance of cabbage and detrimental effects of Cd on growth, based on the research of varietal variation of Cd tolerance and Cd concentration, by using four levels of Cd hydroponic experiments. Meanwhile, the possibility of reducing Cd uptake and accumulation in cabbage by application of chemical regulators was also studied in the investigation. The major results were summarized as follows:
1. Response of different varieties of cabbage to cadmium stress
Pot experiment was conduced to investigate the reaction of31cabbage varieties to Cd toxicity. The results showed that these varieties had significant difference in Cd tolerance, uptake and accumulation when plants grew in the soil contaminated100mg·kg-1Cd. Among the varieties, the inhibition percentage of shoots dry weight varied from-15.4%to91.3%. The detrimental effects of Cd on shoots growth exceeded on roots. The range of Cd concentration in roots was from617to1789mg·kg-1and in shoots from47to183mg·kg-1, which roots exceeded shoots. Based on pot experiment,10varieties screened preliminarily as Cd-tolerance and Cd-sensitivity respectively were planted in a solution with50μmo·L-1 Cd. According to plant tolerance to Cd and the capacity to transport Cd from the roots to the shoots, Chunfeng and lufeng were eventually chosen as the Cd resistant and low Cd transportion variety and the Cd sensitive and high Cd transportion variety, respectively, to investigate the Cd tolerance mechanisms in cabbage.
2. Effects of Cd on growth and Cd uptake of cabbage seedlings
In order to study possible Cd resistance mechanisms in cabbage, several parameters on metal uptake and distribution were compared between the variety Chunfeng (Cd-resistance) and Lufeng (Cd-sensitive).The results showed that low Cd (20μM) exposure for both varieties caused a sharp decline (P<0.05) in the root elongation and the dry weight of plant, with a much more severe responsein the Cd-sensitive varieties. Cd taken up by cabbage plant was mainly distributed in roots compared to shoots, and the variety Chunfeng contained significantly lower Cd concentrations in the leaves and higher Cd ones in roots than Lvfeng. About70-74%and66-68%of the Cd taken up by Lvfeng and Chunfeng, respectively, was transported to shoots. The majority of the Cd accumulated in the leaves, stems and roots of both varieties appeared to be bound on the cell walls. The higher Cd resistance in Chunfeng than in Lufeng may be attributed to higher capacity to limit Cd uptake into shoots and to complex Cd on cell walls.
3. Effects of Cd on photosynthesis, chlorophyll fluorescence and nutrient uptake of cabbage seedlings
Cadmium stress impacted photosynthesis and chlorophyll fluorescence of cabbage plants. Low Cd (20μM) exposure for the Cd-sensitive variety Lufeng caused a sharp decline (P<0.05) in the net photosynthetic rate (Pn), stomatal conductance (Gs), photochemical efficiency of PS Ⅱ (Fv/Fm) and quantum yield of electron flow through PS Ⅱ (OPS Ⅱ) compared to the control. However, these deteriorative effects for the Cd-tolerant variety Chunfeng could be found only at high Cd (50～100μM) levels. At same Cd level, the inhibitions of these parameters were more pronounced in Lufeng than in Chunfeng. Cd stress reduced the concentration of chlorophyll a and chlorophyll b, especially for chlorophyll b, which could be an important factor of photosynthesis inhibition. For these two varieties, Cd stress significantly inhibited the Mn uptake, promoted Zn and S uptake, restrained the transportion of Zn and S (for Lufeng) from roots to shoots, and the decreases of nutrient uptake in lufeng were more obvious than in Chunfeng. Moreover, Cd stress also significantly inhibited Liifeng on Fe and Mg uptake, reduced its Fe and Mg concentration in leaves, but not significant effect on Chunfeng. Cd stress promoted the P uptake by Chunfeng, but opposite results were observed in Lufeng. Therefore, physiologically, the decrease of Mn, Fe, Mg, Zn、Ca and P concentrations in leaves of Lufeng under Cd stress was the key reason for the restraint of leaf photosynthesis, and the decrease of plant growth.
4. Effects of Cd on lipid peroxidation and activities of antioxidant capacity of cabbage seedlings
Excess Cd can cause an increased formation of reactive oxygen species (ROS). However, plants have evolved antioxidantive defence systems to keep ROS under control. The concentration of MDA in Lufeng and Chunfeng plants had not been significantly affected2d and4d, respectively, after Cd treatments, suggesting the antioxidantive defence systems played an importance role, mainly SOD and POD activities significantly increased. Apart from GR activity in leaves of Chunfneg increasing, the SOD, POD, CAT, APX and GR in leaves and roots of two varieties all increased firstly and then decreased with extended Cd exposure of time, but the concentration of MDA significantly increased, which indicated that the increased formation of ROS had exceeded the capacity of antioxidantive defence systems to scavenge ROS, resulting in membrane lipid peroxidation. The activities of antioxidant enzymes in Lufeng plants rised sharply within a short time and then declined significantly compared to that in Chunfeng, to the10th day exposed to Cd, the enzymes all were significantly lower than that in Chunfeng, but the concentration of MDA markedly higher than in Chunfeng, indicating that Lufeng was subjected to a greater degree of oxidative stress. Taken together, Cd induces the formation of ROS, resulting in a lipid peroxidation, which was an important mechanism of Cd toxicity to cabbage plants. The difference of antioxidant capacity might partly explain the Cd resistance of Chunfeng and Lufeng.
5. Role of non-protein thiols and organic acid in Cd tolerance of cabbage
A hydroponic experiment was carried out to examine the role of non-protein thiols and organic acid in Cd tolerance of cabbage. The results showed that compared with the control groups, Cd treatment significantly increased concentrations of non-protein thiol, PCs, citric acids and proline in the leaves and roots of two varieties. The increases were more pronounced in Chunfeng than in Lufeng. Taken together, it is suggested the higher Cd resistance in Chunfeng than in Lvfeng may be attributed to metal binding ligands such as PCs and citric acid, and osmotic adjustment ability. However, the contributions of SH-PCs and citric acid to Cd detoxification might be smaller than that of the cell walls.
6. Role of root cell wall in Cd tolerance of cabbage
Previous research has showed that the cell wall of roots played a important role in cabbage resisting Cd toxicity. To further understand the Cd resistance mechanism in cabbage, changes in cell wall properties of roots under Cd treatments were examined. The results showed that Cd stress evidently increased the concentrations of pectin, hemicellulose I and hemicellulose II in the cell wall of Cd-sensitive variety Liifeng and pectin and hemicellulose II in that of Cd-resistant one Chunfeng. The increase was more pronounced in Lufeng than in Chunfeng. The concentration of the uronic acid in pectin, hemicellulose I and hemicellulose II in Chunfeng and in the pectin and hemicellulose II in Lufeng increased markedly under Cd treatments, but not vary much in hemicellulose I in Lufeng. Chunfeng contained significant higher the uronic acid content than Liifeng, which suggested an important role of the uronic acid in Cd tolerance in cabbage. In addition, among the cell wall polysaccharides in two varieties roots, the Cd content in pectin was the highest, suggesting the majority of the Cd accumulated in root cell walls was tightly bound to the pectin. The Cd concentration in pectin of the variety Chunfeng was significantly higher than that of the variety Lufeng. The results suggested the higher Cd resistance in Chunfeng than in Lvfeng may be attributed to Cd-bound on the uronic acid in pectin.
7. Sodium nitroprusside (as nitric oxide donator) supplementation ameliorates cadmium-toxicity in cabbage
A hydroponic experiment was performed using Cd-sensitive (var. Lufeng) and Cd-tolerant (var. Chunfeng) cabbage seedling to evaluate how different varieties responsed to Cd toxicity in the presence of sodium nitroprusside (SNP), a nitric oxide (NO) donor. Results showed that20-400μM Cd levels inhibited seed germination, and50μM Cd reduced the biomass and chlorophyll concent, with a much more severe response in the Cd-sensitive variety. Cd+SNP treatment restored the Cd-inhibited growth. Exogenous SNP dramatically alleviated Cd toxicity, mainly observed:(a) markedly reduced Cd uptake by plants, obviously diminished Cd accumulation in subcellular fraction of leaves and roots of two varieties,(b) restored the increase of Cd-induced polysaccharides and uronic acid in polysaccharides contents in cell walls of roots,(c) enhanced the activities of antioxidant enzymes and the concentration of antioxidant, whereas decreased the concentrations of malondialdehyde (MDA) and hydrogen peroxide (H2O2), and helped to maintain stability and integrity of membrane system.(d) elevated the increased NP-SH contents, whereas declined slightly in the PCs content, which suggested the GSH may play a important role in protecting cabbage plants from Cd toxicity.(e) promoted nutrients uptake and increased nutrient elements distribution in leavese, and upregulated Cd-induced increase in S content in both varieties, inferring S increment may be involved in NP-SH synthesis;(f) Exogenous SNP reduced the inhibition of Cd on net photosynthetic rate (P n), which might partly be attributed to the NO protection to photosynthetic apparatus such as increasing the chlorophyll content, Fv/Fm, OPS Ⅱ and PSI Ⅰ activity. Taken together, these data suggest that the protection of NO to cabbage seedling against Cd toxicity may be involved in complex signal transduction in plants and cross regulation of physiological and molecular mechanisms.
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