铜处理对菠菜幼苗矿质营养吸收和细胞超微结构的影响
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摘要
土壤重金属积累严重影响植物生长和生态系统平衡,探寻植物对重金属的耐性机理尤为重要.菠菜可能具有一定的耐铜性,但Cu对其矿质元素吸收、细胞超微结构等方面的耐性机理尚不明确.本研究以菠菜幼苗为研究对象,通过盆栽试验,探究不同浓度铜处理对菠菜幼苗生长、矿质元素吸收、叶片细胞超微结构等指标的影响.结果表明:100 mg·L~(-1) CuSO_4处理浓度时,菠菜幼苗根Cu~(2+)积累量小于地上部,其根系生长量增加,地上部生长量稍有下降,继续增加铜处理浓度,植物体各器官生长参数均呈下降趋势.低浓度铜处理时(<400 mg·L~(-1) CuSO_4),菠菜幼苗叶N、K、Ca、Mg、Fe含量增加,P含量减少;根N、P、K含量减少,Ca、Mg、Fe含量增加;叶片细胞内各细胞器清晰可见,基粒片层排列仍较为整齐,叶绿体内外膜完整.高浓度铜处理时(>600 mg·L~(-1) CuSO_4),菠菜幼苗叶N含量增加,P、K、Ca、Mg、Fe含量减少;根N、P、K、Ca、Mg、Fe含量均减少;叶片细胞内叶绿体变圆,叶绿体膜变薄,基质、基粒片层变少,层堆积高度下降,细胞核解体,液泡、细胞壁中有黑色小点分布,可能是大量Cu~(2+)聚集导致细胞内膨压增大所致.低浓度铜处理并未对菠菜幼苗的生长生理特性产生明显的负面影响,而高浓度铜处理并未终止菠菜幼苗的生长.说明菠菜幼苗具有一定的耐铜性.
The accumulation of heavy metals in soil has serious influence on plant growth and ecosystem balance. It is of great importance to explore the mechanism of plant tolerance to heavy me-tals. Although spinach is supposed to have strong Cu tolerance, the effects of Cu on mineral element absorption and cell ultrastructure are still unclear. In this study, the growth of spinach seedlings, the absorption of mineral elements and the ultrastructure of leaf cells were examined in a pot experiment. The results showed that Cu~(2+) accumulation in the root of spinach seedling was less than that in the shoot when CuSO_4 concentration was 100 mg·L~(-1), with root growth being increased and shoot growth being slightly decreased. When copper concentration continued to increase, the growth parameters continuously declined. When the CuSO_(4 c)oncentrations were less than 400 mg·L~(-1), the foliar N, K, Ca, Mg and Fe concentrations of spinach seedling increased, and that of P decreased. The concentrations of N, P and K in roots went down and that of Ca, Mg and Fe went up. All organelles in leaf cells were clearly visible. The basal granule layer was arranged orderly, and the inner and outer membranes of chloroplasts were intact. When the CuSO_4 concentrations exceeded 600 mg·L~(-1), foliar N concentration increased while that of P, K, Ca, Mg and Fe decreased. The concentrations of N, P, K, Ca, Mg and Fe in roots declined. The cell ultrastructure of spinach seedlings substantially changed with the increases of CuSO_4 treated concentrations. The chloroplast in leaf cells became rounder, the chloroplast membrane became thinner, the stroma and basal granule layer became less, and the layer accumulation height decreased. The nucleus was broken up and small black spots were found in vacuoles and cell walls, which might be attributed to the enhancement of intracellular swelling pressure caused by high accumulation of Cu~(2+). In conclusion, low concentration of CuSO_4 had little negative effect on the life activities of spinach seedlings, and the high concentrations of CuSO_4 did not terminate their growth, indicating that spinach seedlings had strong copper resistance.
The accumulation of heavy metals in soil has serious influence on plant growth and ecosystem balance. It is of great importance to explore the mechanism of plant tolerance to heavy me-tals. Although spinach is supposed to have strong Cu tolerance, the effects of Cu on mineral element absorption and cell ultrastructure are still unclear. In this study, the growth of spinach seedlings, the absorption of mineral elements and the ultrastructure of leaf cells were examined in a pot experiment. The results showed that Cu~(2+) accumulation in the root of spinach seedling was less than that in the shoot when CuSO_4 concentration was 100 mg·L~(-1), with root growth being increased and shoot growth being slightly decreased. When copper concentration continued to increase, the growth parameters continuously declined. When the CuSO_(4 c)oncentrations were less than 400 mg·L~(-1), the foliar N, K, Ca, Mg and Fe concentrations of spinach seedling increased, and that of P decreased. The concentrations of N, P and K in roots went down and that of Ca, Mg and Fe went up. All organelles in leaf cells were clearly visible. The basal granule layer was arranged orderly, and the inner and outer membranes of chloroplasts were intact. When the CuSO_4 concentrations exceeded 600 mg·L~(-1), foliar N concentration increased while that of P, K, Ca, Mg and Fe decreased. The concentrations of N, P, K, Ca, Mg and Fe in roots declined. The cell ultrastructure of spinach seedlings substantially changed with the increases of CuSO_4 treated concentrations. The chloroplast in leaf cells became rounder, the chloroplast membrane became thinner, the stroma and basal granule layer became less, and the layer accumulation height decreased. The nucleus was broken up and small black spots were found in vacuoles and cell walls, which might be attributed to the enhancement of intracellular swelling pressure caused by high accumulation of Cu~(2+). In conclusion, low concentration of CuSO_4 had little negative effect on the life activities of spinach seedlings, and the high concentrations of CuSO_4 did not terminate their growth, indicating that spinach seedlings had strong copper resistance.
引文
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[22]Dotaniya ML,Rajendiran S,Coumar MV,et al.Interactive effect of cadmium and zinc on chromium uptake in spinach grown in Vertisol of Central India.International Journal of Environmental Science and Technology,2017,15:441-448
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[24]Bouazizi H,Jouili H,Geitmann A,et al.Copper toxicity in expanding leaves of Phaseolus vulgaris L.:Antioxidant enzyme response and nutrient element uptake.Ecotoxicology&Environmental Safety,2010,73:1304-1308
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[28]Lequeux H,Hermans C,Lutts S,et al.Response to copper excess in Arabidopsis thaliana:Impact on the root system architecture,hormone distribution,lignin accumulation and mineral profile.Plant Physiology and Biochemistry,2010,48:673-682
[29]Xu Q,Qiu H,Chu W,et al.Copper ultrastructural localization,subcellular distribution,and phytotoxicity in Hydrilla verticillata(L.f.)Royle.Environmental Science and Pollution Research International,2013,20:8672-8679
[30]Souza VL,de Almeida AA,Souza Jde S,et al.Altered physiology,cell structure,and gene expression of Theobroma cacao seedlings subjected to Cu toxicity.Environmental Science and Pollution Research International,2014,21:1217-1230
[31]Alia N,Sardar K,Said M,et al.Toxicity and bioaccumulation of heavy metals in spinach(Spinacia oleracea)grown in a controlled environment.International Journal of Environmental Research and Public Health,2015,12:7400-7416
[32]Thounaojam TC,Panda P,Mazumdar P,et al.Excess copper induced oxidative stress and response of antioxidants in rice.Plant Physiology and Biochemistry,2012,53:33-39
[33]Azooz MM,Abouelhamd MF,Alfredan MA.Biphasic effect of copper on growth,proline,lipid peroxidation and antioxidant enzyme activities of wheat(Triticum aestivum cv.Hasaawi)at early growing stage.Australian Journal of Crop Science,2012,6:334-339
[34]Azeez MO,Adesanwo OO,Adepetu JA.Effect of copper(Cu)application on soil available nutrients and uptake.African Journal of Agricultural Research,2015,10:359-364
[35]Chen HC,Jiang JGG.Osmotic adjustment and plant adaptation to environmental changes related to drought and salinity.Environmental Reviews,2010,18:309-319
[36]Sule Guzel,Rabiye Terzi.Exogenous hydrogen peroxide increases dry matter production,mineral content and level of osmotic solutes in young maize leaves and alleviates deleterious effects of copper stress.Botanical Studies,2013,54:26-30
[37]Ali NA,Bernal MP,Ater M.Tolerance and bioaccumulation of copper in Phragmites australis and Zea mays.Plant and Soil,2002,239:103-111
[38]Adrees M,Ali S,Rizwan M,et al.The effect of excess copper on growth and physiology of important food crops:A review.Environmental Science and Pollution Research International,2015,22:8148-8162
[39]Wilkinson SR,Welch RM,Mayland HF,et al.Magnesium in plants:Uptake,distribution,function and utilization by man and animals.Metal Ions in Biological Systems,1990,26:33-56
[40]Janas KM,Zielińska-Tomaszewska J,Rybaczek D,et al.The impact of copper ions on growth,lipid peroxidation,and phenolic compound accumulation and localization in lentil(Lens culinaris Medic.)seedlings.Journal of Plant Physiology,2010,167:270-276
[41]Abdalla FE,El-Saady AM,Safina SA,et al.Response of wheat to magnesium and copper foliar feeding under sandy soil condition.Journal of American Science,2010,6:818-823
[42]Zhang R-R(张然然),Zhang P(张鹏),Du S-T(都韶婷).Oxidative stress-related signals and their regulation under Cd stress.Chinese Journal of Applied Ecology(应用生态学报),2016,27(3):981-992(in chinese)
[43]Monni S,Salemaa M,White C,et al.Copper resistance of Calluna vulgaris originating from the pollution gradient of a Cu-Ni smelter,in southwest Finland.Environmental Pollution,2000,109:211-219
[44]Lou LQ,Shen ZG,Li XD.The copper tolerance mechanisms of Elsholtzia haichowensis,a plant from copperenriched soils.Environmental&Experimental Botany,2004,51:111-120
[45]Terry N,Abadía J.Function of iron in chloroplasts.Journal of Plant Nutrition,1986,9:609-646
[46]Minorsky PV.An heuristic hypothesis of chilling injury in plants:A role for calcium as the primary physiological transducer of injury.Plant,Cell&Environment,2010,8:75-94
[47]Arora R,Palta JP.A loss in the plasma membrane AT-Pase activity and its recovery coincides with incipient freeze-thaw injury and postthaw recovery in onion bulb scale tissue.Plant Physiology,1991,95:846-852
[48]Mozdzeń K,Wanic T,Rut G,et al.Toxic effect on the physiological processes in Pinus sylvestris L.grown under high copper content.Photosynthetica,2016,55:1-8
[49]Mehes-Smith M,Nkongolo K,Cholewa E.Coping mechanisms of plants to metal contaminated soil//Silvern Hall CM,ed.Environmental Change and Sustainability.Berlin,Germany:Spring-Verlag,2013:53-72
[50]Mei X,Li S,Li Q,et al.Sodium chloride salinity reduces Cd uptake by edible amaranth(Amaranthus mangostanus L.)via competition for Ca channels.Ecotoxicology&Environmental Safety,2014,105:59-64
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[2]Zhang X-N(张晓娜),Piao C-L(朴春兰),Dong Y-K(董友魁),et al.Transcriptome analysis of response to heavy metal Cd stress in soybean root.Chinese Journal of Applied Ecology(应用生态学报),2017,28(5):1633-1641(in Chinese)
[3]Guo X-L(郭星亮),Gu J(谷洁),Chen Z-X(陈智学),et al.Effects of heavy metals pollution on soil microbial communities metabolism and soil enzyme activities in coal mining area of Tongchuan,Shaanxi Province of Northwest China.Chinese Journal of Applied Ecology(应用生态学报),2012,23(3):798-806(in Chinese)
[4]Ministry of Environmental Protection,Ministry of Land and Resources of China(国家环境保护部、国土资源部).Bulletin of National Soil Pollution Investigation in2014[EB/OL].(2014-04-17)[2014-04-17]http://www.zhb.gov.cn/gkml/hbb/qt/201404/t20140417_270670.htm(in Chinese)
[5]Dos Santos RW,Schmidt EC,de LFMR,et al.Bioabsorption of cadmium,copper and lead by the red macroalga Gelidium floridanum:Physiological responses and ultrastructure features.Ecotoxicology and Environmental Safety,2014,105:80-89
[6]Yruela I.Copper in plants:Acquisition,transport and interactions.Functional Plant Biology,2009,36:409-430
[7]Cuchiara CC,LarréCF,Peters JA.Copper and zinc interactions:Morphophysiological responses in sweet potato plants(Ipomoea batatas L.).Iheringia,Série Botnica,2017,72:142-149
[8]Liu B-L(刘宝林),Zhao X-M(赵秀敏).Investigation and research of trace elements in spinaches of different places.Journal of Changchun Normal University(Natural Science)(长春师范学院学报:自然科学版),2012,31(9):51-54(in Chinese)
[9]Jia L(贾丽),Guo D-F(郭笃发),Wang L-L(王龙龙).Characteristic of heavy metal accumulation in vegetables in Jinan vegetable fields.Journal of Anhui Agricultural Sciences(安徽农业科学),2015,43(2):117-119(in Chinese)
[10]Fan Y,Li H,Xue ZJ,et al.Accumulation characteristics and potential risk of heavy metals in soil-vegetable system under greenhouse cultivation condition in Northern China.Ecological Engineering,2017,102:367-373
[11]Lisiewska Z,Kmiecik W,Gbczyński P,et al.Amino acid profile of raw and as-eaten products of spinach(Spinacia oleracea L.).Food Chemistry,2011,126:460-465
[12]Becker C,Klaering HP,Kroh LW,et al.Cool-cultivated red leaf lettuce accumulates cyanidin-3-O-(6″-O-malonyl)-glucoside and caffeoylmalic acid.Food Chemistry,2014,146:404-411
[13]Naz S,Anjum MA,Akhtar S.Monitoring of growth,yield,biomass and heavy metals accumulation in spinach grown under different irrigation sources.International Journal of Agriculture and Biology,2016,18:689-697
[14]Singh D,Kumar A.Impact of irrigation using water containing Cu O and Zn O nanoparticles on spinach oleracea grown in soil media.Bulletin of Environmental Contamination and Toxicology,2016,97:548-553
[15]Liu W-Y(刘文英),Zhou F(周凤),Rong T-T(戎婷婷),et al.Effects of heavy metal copper on physiological indexes of spinach.Agriculture and Technology(农业与技术),2016,36(3):1-3(in Chinese)
[16]Chen JR,Peng DL,Shafi M,et al.Effect of copper toxicity on root morphology,ultrastructure,and copper accumulation in moso bamboo(Phyllostachys pubescens).Zeitschrift fur Naturforschung Section C,2014,69:399-406
[17]Han YL,Yuan HY,Huang SZ,et al.Cadmium tolerance and accumulation by two species of Iris.Ecotoxicology,2007,16:557-563
[18]Graaff MAD,Six J,Jastrow JD,et al.Variation in root architecture among switchgrass cultivars impacts root decomposition rates.Soil Biology&Biochemistry,2013,58:198-206
[19]Lu R-K(鲁如坤).Analysis Methods for Soil and Agricultural Chemistry.Nanjing:Hohai University Press,2000:146-147(in Chinese)
[20]Li H-S(李合生).The Experiment Principle and Technique on Plant Physiology and Biochemistry.Beijing:Higher Education Press,2000:121-123(in Chinese)
[21]Israr M,Jewell A,Kumar D,et al.Interactive effects of lead,copper,nickel and zinc on growth,metal uptake and antioxidative metabolism of Sesbania drummondii.Journal of Hazardous Materials,2011,186:1520-1526
[22]Dotaniya ML,Rajendiran S,Coumar MV,et al.Interactive effect of cadmium and zinc on chromium uptake in spinach grown in Vertisol of Central India.International Journal of Environmental Science and Technology,2017,15:441-448
[23]Kumar R,Mehrotra NK,Nautiyal BD,et al.Effect of copper on growth,yield and concentration of Fe,Mn,Zn and Cu in wheat plants(Triticum aestivum L.).Journal of Environmental Biology,2009,30:485-488
[24]Bouazizi H,Jouili H,Geitmann A,et al.Copper toxicity in expanding leaves of Phaseolus vulgaris L.:Antioxidant enzyme response and nutrient element uptake.Ecotoxicology&Environmental Safety,2010,73:1304-1308
[25]Liu WH,Zhao JZ,Ouyang ZY,et al.Impacts of sewage irrigation on heavy metal distribution and contamination in Beijing,China.Environment International,2005,31:805-812
[26]Khan S,Cao Q,Zheng YM,et al.Health risks of heavy metals in contaminated soils and food crops irrigated with wastewater in Beijing,China.Environmental Pollution,2008,152:686-692
[27]Monu A,Bala K,Shweta R,et al.Heavy metal accumulation in vegetables irrigated with water from different sources.Food Chemistry,2008,111:811-815
[28]Lequeux H,Hermans C,Lutts S,et al.Response to copper excess in Arabidopsis thaliana:Impact on the root system architecture,hormone distribution,lignin accumulation and mineral profile.Plant Physiology and Biochemistry,2010,48:673-682
[29]Xu Q,Qiu H,Chu W,et al.Copper ultrastructural localization,subcellular distribution,and phytotoxicity in Hydrilla verticillata(L.f.)Royle.Environmental Science and Pollution Research International,2013,20:8672-8679
[30]Souza VL,de Almeida AA,Souza Jde S,et al.Altered physiology,cell structure,and gene expression of Theobroma cacao seedlings subjected to Cu toxicity.Environmental Science and Pollution Research International,2014,21:1217-1230
[31]Alia N,Sardar K,Said M,et al.Toxicity and bioaccumulation of heavy metals in spinach(Spinacia oleracea)grown in a controlled environment.International Journal of Environmental Research and Public Health,2015,12:7400-7416
[32]Thounaojam TC,Panda P,Mazumdar P,et al.Excess copper induced oxidative stress and response of antioxidants in rice.Plant Physiology and Biochemistry,2012,53:33-39
[33]Azooz MM,Abouelhamd MF,Alfredan MA.Biphasic effect of copper on growth,proline,lipid peroxidation and antioxidant enzyme activities of wheat(Triticum aestivum cv.Hasaawi)at early growing stage.Australian Journal of Crop Science,2012,6:334-339
[34]Azeez MO,Adesanwo OO,Adepetu JA.Effect of copper(Cu)application on soil available nutrients and uptake.African Journal of Agricultural Research,2015,10:359-364
[35]Chen HC,Jiang JGG.Osmotic adjustment and plant adaptation to environmental changes related to drought and salinity.Environmental Reviews,2010,18:309-319
[36]Sule Guzel,Rabiye Terzi.Exogenous hydrogen peroxide increases dry matter production,mineral content and level of osmotic solutes in young maize leaves and alleviates deleterious effects of copper stress.Botanical Studies,2013,54:26-30
[37]Ali NA,Bernal MP,Ater M.Tolerance and bioaccumulation of copper in Phragmites australis and Zea mays.Plant and Soil,2002,239:103-111
[38]Adrees M,Ali S,Rizwan M,et al.The effect of excess copper on growth and physiology of important food crops:A review.Environmental Science and Pollution Research International,2015,22:8148-8162
[39]Wilkinson SR,Welch RM,Mayland HF,et al.Magnesium in plants:Uptake,distribution,function and utilization by man and animals.Metal Ions in Biological Systems,1990,26:33-56
[40]Janas KM,Zielińska-Tomaszewska J,Rybaczek D,et al.The impact of copper ions on growth,lipid peroxidation,and phenolic compound accumulation and localization in lentil(Lens culinaris Medic.)seedlings.Journal of Plant Physiology,2010,167:270-276
[41]Abdalla FE,El-Saady AM,Safina SA,et al.Response of wheat to magnesium and copper foliar feeding under sandy soil condition.Journal of American Science,2010,6:818-823
[42]Zhang R-R(张然然),Zhang P(张鹏),Du S-T(都韶婷).Oxidative stress-related signals and their regulation under Cd stress.Chinese Journal of Applied Ecology(应用生态学报),2016,27(3):981-992(in chinese)
[43]Monni S,Salemaa M,White C,et al.Copper resistance of Calluna vulgaris originating from the pollution gradient of a Cu-Ni smelter,in southwest Finland.Environmental Pollution,2000,109:211-219
[44]Lou LQ,Shen ZG,Li XD.The copper tolerance mechanisms of Elsholtzia haichowensis,a plant from copperenriched soils.Environmental&Experimental Botany,2004,51:111-120
[45]Terry N,Abadía J.Function of iron in chloroplasts.Journal of Plant Nutrition,1986,9:609-646
[46]Minorsky PV.An heuristic hypothesis of chilling injury in plants:A role for calcium as the primary physiological transducer of injury.Plant,Cell&Environment,2010,8:75-94
[47]Arora R,Palta JP.A loss in the plasma membrane AT-Pase activity and its recovery coincides with incipient freeze-thaw injury and postthaw recovery in onion bulb scale tissue.Plant Physiology,1991,95:846-852
[48]Mozdzeń K,Wanic T,Rut G,et al.Toxic effect on the physiological processes in Pinus sylvestris L.grown under high copper content.Photosynthetica,2016,55:1-8
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