硫限制下硒镉胁迫对水稻幼苗的生理特性影响
|
摘要
水稻作为亚洲地区主要粮食作物,在农业生产中占有重要地位。但是随着工业的发展,环境污染日益严重,重金属污染中尤其是镉污染不容忽视。镉为非营养元素,对植物具有明显的毒害作用,而且会通过食物链危及人体健康。如何缓解或者抑制植物镉的毒害,已经引起了人们的广泛关注。本试验以淮稻6号为材料,采用水培的方法,研究了不同硫(S)浓度下(9mg·L-1、720mg·L-1)硒(Se)(5 mg·L-1)和镉(Cd)(1mg·L-1、10mg·L-1)交互胁迫对水稻幼苗的一些生理特性影响,实验中主要研究指标包括生物量、巯基物质含量和硒镉含量。实验研究结果表明：
(1)Cd对水稻幼苗各部分生长状况抑制能力大小不同,其中Cd对根长的抑制作用明显大于对茎长的抑制；
(2)在硒镉实验中,在无Se条件下,随着Cd浓度的增加,Cd对水稻幼苗的生长毒性不断增大,在加入一定浓度的Se后,Cd的毒性作用得到缓解；
(3)在不同浓度S处理和Se、Cd交互实验下,随着S浓度增大,Cd对水稻幼苗毒性作用减轻；但高浓度的S处理反而对Cd与Se交互实验中的Cd毒性产生协同作用；
(4)高浓度的S处理可以抑制水稻幼苗对Se的吸收；
(5)巯基物质含量随着胁迫时间的增加呈先上升后下降的趋势。在Se、Cd交互实验中,低浓度S处理可以促进水稻幼苗中的GSH和PCs含量,从而提高植株体内重金属和PC的络合作用,缓减Cd对水稻幼苗的毒害。
Rice as main food in Asia areas plays an important role in daily agricultural production of human being. However, with the industrial development, environmental pollution is being more and more serious, such as heavy metals (especially cadmium) pollution. Cadmium, a non-nutrients element, has significant toxic effects on plant, which will endanger man's health through the food chain. How to ease or inhibit the cadmium poisoning in plant, has attracted widespread attention.
To better understand the interactions between sulfate and selenium in plant response to cadmium stress, changes in plant growth and the content of non-protein thiols (NPT, GSH, PC) were investigated in No.6 Huai rice seedlings grown hydroponically with Cd, with or without applications of sulfate and selenium. Results of the study are as followed:
(1) The inhibition effects on different parts of rice are various. Results indicated that the inhibition of Cd on the root length was significantly greater than the inhibition of stem length;
(2) In the absence of Se treatment, Cd toxicity on the growth of rice seedlings increased with the enhancement of Cd content in plant. The addition of Se treatment could significantly increased the plant growth;
(3) In the Cd, Se and S combination experiments, Cd toxicity to rice seedlings reduced with alleviation content of S in plant. However, A combination of Se(5 mg·L-1)-S(720 mg·L-1) treatment led to a decrease of rice plants mass;
(4) High concentrations of S treatment can inhibit the absorption of Se in rice seedlings;
(5) The content of non-protein thiols in rice seedlings fist increased and then decreased during the exposure time. In the Cd, Se and S combination experiments, the contents of GSH and PCs enhanced with low concentration of S addition in plant, which promoted the complexation between Cd and non-protein thiols for decreasing the Cd toxicity in rice growth.
(1)Cd对水稻幼苗各部分生长状况抑制能力大小不同,其中Cd对根长的抑制作用明显大于对茎长的抑制；
(2)在硒镉实验中,在无Se条件下,随着Cd浓度的增加,Cd对水稻幼苗的生长毒性不断增大,在加入一定浓度的Se后,Cd的毒性作用得到缓解；
(3)在不同浓度S处理和Se、Cd交互实验下,随着S浓度增大,Cd对水稻幼苗毒性作用减轻；但高浓度的S处理反而对Cd与Se交互实验中的Cd毒性产生协同作用；
(4)高浓度的S处理可以抑制水稻幼苗对Se的吸收；
(5)巯基物质含量随着胁迫时间的增加呈先上升后下降的趋势。在Se、Cd交互实验中,低浓度S处理可以促进水稻幼苗中的GSH和PCs含量,从而提高植株体内重金属和PC的络合作用,缓减Cd对水稻幼苗的毒害。
Rice as main food in Asia areas plays an important role in daily agricultural production of human being. However, with the industrial development, environmental pollution is being more and more serious, such as heavy metals (especially cadmium) pollution. Cadmium, a non-nutrients element, has significant toxic effects on plant, which will endanger man's health through the food chain. How to ease or inhibit the cadmium poisoning in plant, has attracted widespread attention.
To better understand the interactions between sulfate and selenium in plant response to cadmium stress, changes in plant growth and the content of non-protein thiols (NPT, GSH, PC) were investigated in No.6 Huai rice seedlings grown hydroponically with Cd, with or without applications of sulfate and selenium. Results of the study are as followed:
(1) The inhibition effects on different parts of rice are various. Results indicated that the inhibition of Cd on the root length was significantly greater than the inhibition of stem length;
(2) In the absence of Se treatment, Cd toxicity on the growth of rice seedlings increased with the enhancement of Cd content in plant. The addition of Se treatment could significantly increased the plant growth;
(3) In the Cd, Se and S combination experiments, Cd toxicity to rice seedlings reduced with alleviation content of S in plant. However, A combination of Se(5 mg·L-1)-S(720 mg·L-1) treatment led to a decrease of rice plants mass;
(4) High concentrations of S treatment can inhibit the absorption of Se in rice seedlings;
(5) The content of non-protein thiols in rice seedlings fist increased and then decreased during the exposure time. In the Cd, Se and S combination experiments, the contents of GSH and PCs enhanced with low concentration of S addition in plant, which promoted the complexation between Cd and non-protein thiols for decreasing the Cd toxicity in rice growth.
引文
[1]Zhu J M, Zhang B S. Distribution of selenium in a mini-landscape of Yutangba, Enshi, Hubei Province, China [J]. Applied Geochemistry,2001,16:1333-1344
[2]姚林波,高振敏,杨竹森,等.渔塘坝硒矿床富硒硅质岩的成因[J].中国科学(D辑),2002,32(1)：54-63
[3]张光弟,葛晓立,张绮玲,等.湖北恩施硒中毒区土壤硒的分布及其控制因素[J].中国地质,2001,28(9)：37-41
[4]Lin K F, Xiang Y L. Metallic Elements in Hair as a Biomarker of Human Exposure to Environmental Pollution [J]. CRC Critical Reveny in Plant Science,18(3):417-428
[5]林匡飞,徐小清.As污染区农民头发中As含量与环境中As含量的关系[J].中国环境科学,2001,21(5)：440-444
[6]林匡飞.硒和锗在土壤—植物系统中的生态毒理效应研究.中国科学院博士学位研究生学位论文,2003：191-212
[7]吴志刚,李郧,陈学敏.硒镉联合注射染毒法的方法学研究[J].中国公共卫生,2002,18(5)：531-532
[8]Sasakara C, Kazuo T S. Biological interaction between transition metals (Ag, Cd, and Hg) Selenide/sulfide and selenoprotein [J]. Journal of Inorganic Biochem.,1998,71:159-162
[9]Magos L, Webb M. The interaction of selenium with cadmium and mercury [J]. CRC Critical Reveny Toxicol,1980(8):1-42
[10]Nishiyama S, Nakamura K, Konish Y. Effect of Selenium on blood pressure, urinary sodium excretion and plasma aldosterone in cadmium treated male rats [J]. Toxicol, 1987,59:365-370
[11]彭双清,虞昊,刘世杰.硒对镉致脂质过氧化与金属硫化蛋白诱导合成的影响[J].卫生毒理学杂志,2003,17(3)：155-158
[12]王爱国,陈学敏,鲁文清,等.低硒高镉对大鼠组织中谷胱甘肽过氧化酶活性影响的动态观察[J].中国地方病学杂志,1995,14(6)：337-340
[13]王宗元,王捍东,史德浩,等.镉中毒解毒机理的研究—硒化合物的预防和治疗作用初探[J].畜牧兽医学报,1993,24(4)：366-373
[14]段柳艳,高腾云,傅彤,等.硒的营养作用及在反刍动物中的应用[J].2009(7)：39-40,48
[15]罗世炜,史丽英,吴永尧.植物硒研究进展[J].生物学教学,2008,33(1)：2-3
[16]魏筱红,魏泽义.镉的毒性及其危害[J].公共卫生与预防医学,2007,18(4)：4-46
[17]Wu X, Jin T, Wang Z, et al. Urinary calcium as biomarker of renal dysfunction in a general population exposed to Cadmium [J]. Occup Environ Vied,2001,43:898-904
[18]秦光蔚,梁永超,周祥,等.硫对油菜产量和抗逆性的影响[J].土壤肥料,2001,1：36-39
[19]Wu X, Jin T, Wang Z, et al. Urinary calcium as biomarker of renal dysfunction in a general populat ion exposed to Cadmium [J]. Journal of Occupational and Environmental Medicine,2001,43:898-904
[20]尚庆茂,高丽红,李式军.硒素营养对水培生菜品质的影响[J].中国农业大学学报,1998,3(3)：67-71
[21]马友华,丁瑞兴,张继榛,等.植物体内硒和硫的相互作用[J].植物生理学通讯,2001,37(2)：161-166
[22]张驰,吴永尧,彭振坤,等.油菜苗期对硒的生物富集分布[J].河南农业科学,2005(9)：29-32
[23]Vallee B L, Ulmer D D. Biochermical effects of mercury, cadmium and lead [J]. Annual Reveny Biochemistry,1972,41:92-108
[24]杨志新,刘树庆.土壤重金属复合污染对油菜生长的影响[J].河北农业大学学报,2000,23(3)：27-30
[25]王涛,郭智,奥岩松.镉对龙葵幼苗生长的影响及镉富集特性研究[J].上海交通大学学报(农业科学版),2009,27(3)：200-205
[26]徐伟慧,王志刚.镉对菜豆种子萌发及蛋白质含量的影响[J].安徽农业科学,2009,37(18)：8430-8431,8460
[27]Grill E, Winnacker E L, Zenk M H. Phytochelatins, a class of heavy-metal-binding peptides from plants, are functionally analogous to metallothioneins [J]. Proceedings of the National Academy of Sciences,1987,84:439-443
[28]彭诚.硒对镉协迫下白菜幼苗生理特性的影响[J].湖北民族学院学报(自然科学版),2008,26(1)：97-99
[29]王立新,张海芸,郁建锋,等.硒对汞、镉复合污染下豌豆幼苗生长的影响[J].常熟理工学院学报(自然科学),2009,23(4)：71-74
[30]Stobart A K, GriffithsW T, Ameen-Bukharil, et al. The effect of Cd2+on the biosynthesis chlorophyll in leaves of barley [J]. Physiol Plant,1985,63:293-298
[31]谷巍,施国新,杜开和,等.汞、镉复合污染对轮叶狐尾藻的毒害影响[J].南京师大学报(自然科学版),2001,24(3)：75-79
[32]廖琳,胡晓荣,李晖,等.生态环境中镉对生物体毒性作用机理及硒对该毒性拮抗作用的研究进展[J].四川环境,2002,21(2)：21-24
[33]Rana S V. Protective effects of GSH, VitaminE and selenium on lipid peroxidation in cadmium-fed rats [J]. Biological Trace Element Re-search,1996,51:161-168
[34]Jamall I S, Smith J C. Effects of cadmium on glutathione perpoxidase, superoxide dismutase and lipidperoxidation in the rat heart:a possible mechanism of cadmium cardiotoxicity [J]. Toxjicol Applpharmecol Toxicol,1987,60:355-358
[35]张华,史红文,邱喜阳,等.镉-硫-硒复合作用下硒在土壤-蔬菜系统中的迁移分配研究[J].湖南科技大学学报(自然科学版),2009,24(2)：112-115
[36]孙新,杨志敏,徐朗莱.缺硫条件下油菜对镉毒害的敏感性[J].南京农业大学学报,2003,26(4)：56-59
[37]孙新,杨志敏,徐朗莱.镉诱导油菜叶片氧化胁迫及硫化物的络合作用[J].西北植物学报,2003,23(9)：1506-1511
[38]刘丽光,任长林.硫与硒在心肌损害中相关关系的实验研究[J].职业卫生与应急救援,1997,15(2)：73-74
[39]Reese R N, White C A, Winge D R. Long-distance transport of reduced sulphur in spruce (Picea abies L.) [J]. Journal of Experimental Botany,1992,43:1243-1250
[40]黄峙,郑文杰,向军俭,等.硒硫比值对钝顶螺旋藻有机化硒的影响及藻体中硒的形态、价态构成[J].海洋科学,2002,26(5)：60-62
[41]李登超,朱祝军,徐志豪.不同硫水平下硒对小白菜生长及养分含量的影响[J].浙江大学学报(农业与生命科学版),2003,29(4)：402-406
[42]马友华,丁瑞兴,张继榛等.硒和硫配施对烟草叶绿素及保护酶活性的影响[J].南京农业大学学报,1999,22(4)：109-111
[43]Ryan J A, Pahren H R, Lucas J B. Controlling cadmium in human chain:review and rationale based on health effect [J]. Environmental Research,1982,28:251-302
[44]Nicholson F A, Jones K C. Effect of phosphate fertilizers and atmospheric deposition on long-time changes in the cadmium content of soil and crop [J]. Environmental Science and Technology,1994,28:2170-2175
[45]Mebarg A A. The role of the plasmalemima in metal tolerance in angiosperms [J]. Physoilogra Plantarum,1993,88:191-198
[46]Ma J F, Hiradate S, Matsumoto H. High aluminum resistance in buckwheat oxalic acid detoxifies aluminum internally [J]. Plant physiol.,1998,117:753-759
[47]Sanita di Toppi L, Gabbrielli R. Response to cadmium in higher plants [J]. Journal of Experimental Botany,1999,41:105-130
[48]Zenk M H. Heavy metal detoxification in higher plant:a review [J]. Gene,1996,179: 21-30
[49]Cobbett C S. Phytochelatin and their roles in heavy metal detoxification [J]. Plant Physiol.,2000,123:825-832
[50]Grill E, Winnacker E L, Zenk M H. Phytochelatins:the principal heavy-metal complexing peptides of higher plants [J]. Science,1985,230:674-676
[51]Maaccnza B, Delana S, Pintore M. The complexes of cadmium with phytochelatins:a quantum mechanicsstudy.In:Wenzel WW. Adriano on the Biogeochemistry of Trace Elements [J]. Austria.International Society for Trance Element Research,1999:884-885
[52]Cobbett C S. A family of phytochelatin synthase genes from plant fungal and animal species [J]. Trends in Plant Science.,1999,4(9):335-336
[53]Vatamaniuk O K, Mari S, Lu Y P, et al. At PCSI, a phytochelatin synthase from Arabidopsis:Isolation and in vitro reconstitution [J]. Proceedings of the National Academy of Sciences,1999,96:7110-7115
[54]Ha S B, Smith A P, Howden R. Phytochelatin symthase gene from Arabtdopsts and the yeast Schizosaccharomyces pomber [J]. Plant Cell,1999,11:1153-1163
[55]Yong L Z, Pilon-Smits E A H, Tarun A S, et al. Cadmium tolerance and accumulation in Indian mustard is enhanced by overexpressing y-glutamylcysteine synthetase [J]. Plant Physiol.,1999,121:1169-1177
[56]Schafer H J, Haag K A, Rausch T. Cdna cloning and expression analysis of genes encoding GSH synthesis in roots of the heavy metal accumulator Brassica Juncea L. evidence or Cd-induction of a putative mitochondrial γ-glutamylcysteine synthetase isoform [J]. Plant Molecular Biology,1998,37:87-97
[57]Milla M A R, Maurer A, Huete A R, et al. Glutathione peroxidase genes in A rabidopsis are ubiquitous and regulated by abiotic stresses through diverse signaling pathways [J]. The plant Journal,2003,36(5):602-615
[58]周启星,程云,张倩茹,等.复合污染生态毒理效应的定量关系分析[J].中国科学C(辑),2003,33(6)：566-573
[59]郑振华,周培疆,吴振斌.复合污染研究的新进展[J].应用生态学报,2001,12(3)：469-473
[60]Silva E, Rajapakse N, Kortenkamp A. Something from "Nothing"-Eight weak Estrogenic chemicals combined at concentrations below NOECs produce significant mixture effects [J]. Environmental Science and Technology,2002,36:1751-1756
[61]Eggen R I L, Behra R, Patricia B H, et al. Challenges in ecotoxicology [J]. Environmental Science and Technology,2004:58A-64A
[62]Cary, E E. Effect of selenium and cadmium additions to soil on their concentration in lettuce and wheat [J]. Agronomy journal,1981,73:703-708
[63]Shanker K, Mishra S, Srivastava S, et al. Studies on Cd-Se interaction with reference to the uptake and translocation of cadmium in kidney bean [J]. Chemical Speciation and Bioavailability,1995,7(3):97-100
[64]He P P, Lv X Z, Wang G Y. Effects of Se and Zn supplementation on the antagonism against Pb and Cd in vegetables [J]. Environmental International,2004,30:67-172
[65]陈平,余土元,陈惠阳,等.硒对镉胁迫下水稻幼苗生长及生理特征的影响[J].广西植物,2002,22(3)：277-282
[66]陈平,吴秀峰,张伟峰,等.硒对镉胁迫下水稻幼苗叶片元素含量的影响[J].中国生态农业学报,2006,14(3)：277-282
[67]Leustek T, Saito K. Sulfate transport and assimilation in plants [J]. Plant Physiol,1999, 120:637-643
[68]Di Toppi S D, Gabbrielli R. Response to cadmium in higher plants [J]. Journal of Experimental Botany,1999,41:105-130
[69]Heiss S, Schafer H J, Haag-Kerwer A, et al. Cloning sulfur assimilation genes of Brassica juncea L:Cadmium differentially affects the expression of a putative low-affinity sulfate transport and isoforms of ATP sulfurylase and APS reductase [J]. Plant Mol Boil,1999,39:847-857
[70]Lee S, Leustek T. The affect of cadmium on sulfate assimilation enzymes in Brassica juncea [J]. Plant Science,1999,141:201-207
[71]Clemens S, Palmgren M G, Kramer U. A long way ahead:understanding and engineering plant metal accumulation [J]. Trends Plant Science,2002,7:309-315
[72]Bailey F C, Knight A W, Ogle R S, et al. Effect of sulfate level on selenium uptake by Ruppia maritime [J]. Chemosphere,1995,30(3):579-591
[73]Rauser W E. Structure and function of metal chelators produced by plants:the case for organic acids, amino acids, phytin, and metallothioneins [J]. Cell Biophys,1999,31: 19-48
[74]陈同斌,黄泽春,黄宇营,等.砷超富集植物中元素的微区分布及其与砷富集的关系[J].科学通报,2003,48(11)：1163-1168
[75]Howe J A, Loeppert R H, Derose V J, et al. Localization and speciation of chromium in subterranean clover using XRF, XANES, and EPR spectroscopy [J]. Environmental Science and Technology,2003,37(18):4091-4097
[76]S Rama Devi, et al. Copper toxicity in Ceratophyllum demersum L. (Coontail), a free floating macrophyte:Response of antioxidant enzymes and antioxidants [J]. Plant Science,1998,138(2):157-165
[77]赵旭东,魏东芝,万群,等.谷胱甘肽的简便测定法[J].药物分析杂志,2000,20(1)：34-37
[78]Poonam B, Ashish K S, Snigdha U, et al. Phytochelatin plays a role in UV-B tolerance in N2-fixing cyanobacterium Anabaena doliolum [J]. Journal of Plant Physiology,2005, 16(11):1220-1225
[2]姚林波,高振敏,杨竹森,等.渔塘坝硒矿床富硒硅质岩的成因[J].中国科学(D辑),2002,32(1)：54-63
[3]张光弟,葛晓立,张绮玲,等.湖北恩施硒中毒区土壤硒的分布及其控制因素[J].中国地质,2001,28(9)：37-41
[4]Lin K F, Xiang Y L. Metallic Elements in Hair as a Biomarker of Human Exposure to Environmental Pollution [J]. CRC Critical Reveny in Plant Science,18(3):417-428
[5]林匡飞,徐小清.As污染区农民头发中As含量与环境中As含量的关系[J].中国环境科学,2001,21(5)：440-444
[6]林匡飞.硒和锗在土壤—植物系统中的生态毒理效应研究.中国科学院博士学位研究生学位论文,2003：191-212
[7]吴志刚,李郧,陈学敏.硒镉联合注射染毒法的方法学研究[J].中国公共卫生,2002,18(5)：531-532
[8]Sasakara C, Kazuo T S. Biological interaction between transition metals (Ag, Cd, and Hg) Selenide/sulfide and selenoprotein [J]. Journal of Inorganic Biochem.,1998,71:159-162
[9]Magos L, Webb M. The interaction of selenium with cadmium and mercury [J]. CRC Critical Reveny Toxicol,1980(8):1-42
[10]Nishiyama S, Nakamura K, Konish Y. Effect of Selenium on blood pressure, urinary sodium excretion and plasma aldosterone in cadmium treated male rats [J]. Toxicol, 1987,59:365-370
[11]彭双清,虞昊,刘世杰.硒对镉致脂质过氧化与金属硫化蛋白诱导合成的影响[J].卫生毒理学杂志,2003,17(3)：155-158
[12]王爱国,陈学敏,鲁文清,等.低硒高镉对大鼠组织中谷胱甘肽过氧化酶活性影响的动态观察[J].中国地方病学杂志,1995,14(6)：337-340
[13]王宗元,王捍东,史德浩,等.镉中毒解毒机理的研究—硒化合物的预防和治疗作用初探[J].畜牧兽医学报,1993,24(4)：366-373
[14]段柳艳,高腾云,傅彤,等.硒的营养作用及在反刍动物中的应用[J].2009(7)：39-40,48
[15]罗世炜,史丽英,吴永尧.植物硒研究进展[J].生物学教学,2008,33(1)：2-3
[16]魏筱红,魏泽义.镉的毒性及其危害[J].公共卫生与预防医学,2007,18(4)：4-46
[17]Wu X, Jin T, Wang Z, et al. Urinary calcium as biomarker of renal dysfunction in a general population exposed to Cadmium [J]. Occup Environ Vied,2001,43:898-904
[18]秦光蔚,梁永超,周祥,等.硫对油菜产量和抗逆性的影响[J].土壤肥料,2001,1：36-39
[19]Wu X, Jin T, Wang Z, et al. Urinary calcium as biomarker of renal dysfunction in a general populat ion exposed to Cadmium [J]. Journal of Occupational and Environmental Medicine,2001,43:898-904
[20]尚庆茂,高丽红,李式军.硒素营养对水培生菜品质的影响[J].中国农业大学学报,1998,3(3)：67-71
[21]马友华,丁瑞兴,张继榛,等.植物体内硒和硫的相互作用[J].植物生理学通讯,2001,37(2)：161-166
[22]张驰,吴永尧,彭振坤,等.油菜苗期对硒的生物富集分布[J].河南农业科学,2005(9)：29-32
[23]Vallee B L, Ulmer D D. Biochermical effects of mercury, cadmium and lead [J]. Annual Reveny Biochemistry,1972,41:92-108
[24]杨志新,刘树庆.土壤重金属复合污染对油菜生长的影响[J].河北农业大学学报,2000,23(3)：27-30
[25]王涛,郭智,奥岩松.镉对龙葵幼苗生长的影响及镉富集特性研究[J].上海交通大学学报(农业科学版),2009,27(3)：200-205
[26]徐伟慧,王志刚.镉对菜豆种子萌发及蛋白质含量的影响[J].安徽农业科学,2009,37(18)：8430-8431,8460
[27]Grill E, Winnacker E L, Zenk M H. Phytochelatins, a class of heavy-metal-binding peptides from plants, are functionally analogous to metallothioneins [J]. Proceedings of the National Academy of Sciences,1987,84:439-443
[28]彭诚.硒对镉协迫下白菜幼苗生理特性的影响[J].湖北民族学院学报(自然科学版),2008,26(1)：97-99
[29]王立新,张海芸,郁建锋,等.硒对汞、镉复合污染下豌豆幼苗生长的影响[J].常熟理工学院学报(自然科学),2009,23(4)：71-74
[30]Stobart A K, GriffithsW T, Ameen-Bukharil, et al. The effect of Cd2+on the biosynthesis chlorophyll in leaves of barley [J]. Physiol Plant,1985,63:293-298
[31]谷巍,施国新,杜开和,等.汞、镉复合污染对轮叶狐尾藻的毒害影响[J].南京师大学报(自然科学版),2001,24(3)：75-79
[32]廖琳,胡晓荣,李晖,等.生态环境中镉对生物体毒性作用机理及硒对该毒性拮抗作用的研究进展[J].四川环境,2002,21(2)：21-24
[33]Rana S V. Protective effects of GSH, VitaminE and selenium on lipid peroxidation in cadmium-fed rats [J]. Biological Trace Element Re-search,1996,51:161-168
[34]Jamall I S, Smith J C. Effects of cadmium on glutathione perpoxidase, superoxide dismutase and lipidperoxidation in the rat heart:a possible mechanism of cadmium cardiotoxicity [J]. Toxjicol Applpharmecol Toxicol,1987,60:355-358
[35]张华,史红文,邱喜阳,等.镉-硫-硒复合作用下硒在土壤-蔬菜系统中的迁移分配研究[J].湖南科技大学学报(自然科学版),2009,24(2)：112-115
[36]孙新,杨志敏,徐朗莱.缺硫条件下油菜对镉毒害的敏感性[J].南京农业大学学报,2003,26(4)：56-59
[37]孙新,杨志敏,徐朗莱.镉诱导油菜叶片氧化胁迫及硫化物的络合作用[J].西北植物学报,2003,23(9)：1506-1511
[38]刘丽光,任长林.硫与硒在心肌损害中相关关系的实验研究[J].职业卫生与应急救援,1997,15(2)：73-74
[39]Reese R N, White C A, Winge D R. Long-distance transport of reduced sulphur in spruce (Picea abies L.) [J]. Journal of Experimental Botany,1992,43:1243-1250
[40]黄峙,郑文杰,向军俭,等.硒硫比值对钝顶螺旋藻有机化硒的影响及藻体中硒的形态、价态构成[J].海洋科学,2002,26(5)：60-62
[41]李登超,朱祝军,徐志豪.不同硫水平下硒对小白菜生长及养分含量的影响[J].浙江大学学报(农业与生命科学版),2003,29(4)：402-406
[42]马友华,丁瑞兴,张继榛等.硒和硫配施对烟草叶绿素及保护酶活性的影响[J].南京农业大学学报,1999,22(4)：109-111
[43]Ryan J A, Pahren H R, Lucas J B. Controlling cadmium in human chain:review and rationale based on health effect [J]. Environmental Research,1982,28:251-302
[44]Nicholson F A, Jones K C. Effect of phosphate fertilizers and atmospheric deposition on long-time changes in the cadmium content of soil and crop [J]. Environmental Science and Technology,1994,28:2170-2175
[45]Mebarg A A. The role of the plasmalemima in metal tolerance in angiosperms [J]. Physoilogra Plantarum,1993,88:191-198
[46]Ma J F, Hiradate S, Matsumoto H. High aluminum resistance in buckwheat oxalic acid detoxifies aluminum internally [J]. Plant physiol.,1998,117:753-759
[47]Sanita di Toppi L, Gabbrielli R. Response to cadmium in higher plants [J]. Journal of Experimental Botany,1999,41:105-130
[48]Zenk M H. Heavy metal detoxification in higher plant:a review [J]. Gene,1996,179: 21-30
[49]Cobbett C S. Phytochelatin and their roles in heavy metal detoxification [J]. Plant Physiol.,2000,123:825-832
[50]Grill E, Winnacker E L, Zenk M H. Phytochelatins:the principal heavy-metal complexing peptides of higher plants [J]. Science,1985,230:674-676
[51]Maaccnza B, Delana S, Pintore M. The complexes of cadmium with phytochelatins:a quantum mechanicsstudy.In:Wenzel WW. Adriano on the Biogeochemistry of Trace Elements [J]. Austria.International Society for Trance Element Research,1999:884-885
[52]Cobbett C S. A family of phytochelatin synthase genes from plant fungal and animal species [J]. Trends in Plant Science.,1999,4(9):335-336
[53]Vatamaniuk O K, Mari S, Lu Y P, et al. At PCSI, a phytochelatin synthase from Arabidopsis:Isolation and in vitro reconstitution [J]. Proceedings of the National Academy of Sciences,1999,96:7110-7115
[54]Ha S B, Smith A P, Howden R. Phytochelatin symthase gene from Arabtdopsts and the yeast Schizosaccharomyces pomber [J]. Plant Cell,1999,11:1153-1163
[55]Yong L Z, Pilon-Smits E A H, Tarun A S, et al. Cadmium tolerance and accumulation in Indian mustard is enhanced by overexpressing y-glutamylcysteine synthetase [J]. Plant Physiol.,1999,121:1169-1177
[56]Schafer H J, Haag K A, Rausch T. Cdna cloning and expression analysis of genes encoding GSH synthesis in roots of the heavy metal accumulator Brassica Juncea L. evidence or Cd-induction of a putative mitochondrial γ-glutamylcysteine synthetase isoform [J]. Plant Molecular Biology,1998,37:87-97
[57]Milla M A R, Maurer A, Huete A R, et al. Glutathione peroxidase genes in A rabidopsis are ubiquitous and regulated by abiotic stresses through diverse signaling pathways [J]. The plant Journal,2003,36(5):602-615
[58]周启星,程云,张倩茹,等.复合污染生态毒理效应的定量关系分析[J].中国科学C(辑),2003,33(6)：566-573
[59]郑振华,周培疆,吴振斌.复合污染研究的新进展[J].应用生态学报,2001,12(3)：469-473
[60]Silva E, Rajapakse N, Kortenkamp A. Something from "Nothing"-Eight weak Estrogenic chemicals combined at concentrations below NOECs produce significant mixture effects [J]. Environmental Science and Technology,2002,36:1751-1756
[61]Eggen R I L, Behra R, Patricia B H, et al. Challenges in ecotoxicology [J]. Environmental Science and Technology,2004:58A-64A
[62]Cary, E E. Effect of selenium and cadmium additions to soil on their concentration in lettuce and wheat [J]. Agronomy journal,1981,73:703-708
[63]Shanker K, Mishra S, Srivastava S, et al. Studies on Cd-Se interaction with reference to the uptake and translocation of cadmium in kidney bean [J]. Chemical Speciation and Bioavailability,1995,7(3):97-100
[64]He P P, Lv X Z, Wang G Y. Effects of Se and Zn supplementation on the antagonism against Pb and Cd in vegetables [J]. Environmental International,2004,30:67-172
[65]陈平,余土元,陈惠阳,等.硒对镉胁迫下水稻幼苗生长及生理特征的影响[J].广西植物,2002,22(3)：277-282
[66]陈平,吴秀峰,张伟峰,等.硒对镉胁迫下水稻幼苗叶片元素含量的影响[J].中国生态农业学报,2006,14(3)：277-282
[67]Leustek T, Saito K. Sulfate transport and assimilation in plants [J]. Plant Physiol,1999, 120:637-643
[68]Di Toppi S D, Gabbrielli R. Response to cadmium in higher plants [J]. Journal of Experimental Botany,1999,41:105-130
[69]Heiss S, Schafer H J, Haag-Kerwer A, et al. Cloning sulfur assimilation genes of Brassica juncea L:Cadmium differentially affects the expression of a putative low-affinity sulfate transport and isoforms of ATP sulfurylase and APS reductase [J]. Plant Mol Boil,1999,39:847-857
[70]Lee S, Leustek T. The affect of cadmium on sulfate assimilation enzymes in Brassica juncea [J]. Plant Science,1999,141:201-207
[71]Clemens S, Palmgren M G, Kramer U. A long way ahead:understanding and engineering plant metal accumulation [J]. Trends Plant Science,2002,7:309-315
[72]Bailey F C, Knight A W, Ogle R S, et al. Effect of sulfate level on selenium uptake by Ruppia maritime [J]. Chemosphere,1995,30(3):579-591
[73]Rauser W E. Structure and function of metal chelators produced by plants:the case for organic acids, amino acids, phytin, and metallothioneins [J]. Cell Biophys,1999,31: 19-48
[74]陈同斌,黄泽春,黄宇营,等.砷超富集植物中元素的微区分布及其与砷富集的关系[J].科学通报,2003,48(11)：1163-1168
[75]Howe J A, Loeppert R H, Derose V J, et al. Localization and speciation of chromium in subterranean clover using XRF, XANES, and EPR spectroscopy [J]. Environmental Science and Technology,2003,37(18):4091-4097
[76]S Rama Devi, et al. Copper toxicity in Ceratophyllum demersum L. (Coontail), a free floating macrophyte:Response of antioxidant enzymes and antioxidants [J]. Plant Science,1998,138(2):157-165
[77]赵旭东,魏东芝,万群,等.谷胱甘肽的简便测定法[J].药物分析杂志,2000,20(1)：34-37
[78]Poonam B, Ashish K S, Snigdha U, et al. Phytochelatin plays a role in UV-B tolerance in N2-fixing cyanobacterium Anabaena doliolum [J]. Journal of Plant Physiology,2005, 16(11):1220-1225