高锰胁迫下空心莲子草的生理生化特性和草甘膦耐性研究
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摘要
空心莲子草Alternanthera philoxeroides(Mart．)Griseb．是我国危害最为严重的外来入侵杂草之一。鉴于我国高锰胁迫趋势不断加剧，本论文采用水培方法以水生生态型空心莲子草为试验材料，运用ICP-MS、同位素示踪等技术手段，较系统地研究了高锰胁迫对空心莲子草生长与繁殖的影响，高锰胁迫下空心莲子草体内离子吸收、光合与抗氧化酶系统响应，和空心莲子草对草甘膦的耐药性以及草甘膦在杂草体内的吸收、传导与代谢。另外，还针对该草的治理探讨了草甘膦在空心莲子草植株上的沉积规律。取得的初步研究结果如下：
(1)温室条件下空心莲子草生物量调查结果表明，锰浓度在0．31～2．45 mM范围内，培养100 d后空心莲子草茎叶生物量显著高于常规浓度锰(0．0091 mM，对照)培养的植株，锰浓度超过4．90 mM后，空心莲子草生物量随锰浓度进一步提高而下降。次高锰(0．31 mM)条件下培养40、120、365 d后从扦插主根茎上长出的新根茎数是对照的2．3～2．6倍；地下组织生物量比对照提高了24．1％～32．2％；一年的茎叶生物量之和比对照(17．5 g／株)提高了18．4％，均存在显著差异。高锰(2．45 mM)条件下培养40、120、365 d后新根茎数是对照的2．4～3．0倍；培养40 d时地下组织生物量只有对照的82．2％，尔后逐步增加，培养365 d时比对照增加了25．1％；但一年的茎叶生物量之和显著下降，只有对照的71．8％。本实验结果表明，空心莲子草的入侵能力与土壤中有效锰含量密切相关，次高锰促进了空心莲子草生长和地下根茎形成；高锰有利于地下根茎形成，但抑制了地上部生长。
(2)光合速率与荧光参数测定结果表明，不同浓度锰处理后120 d，空心莲子草的光饱和点不受影响，均在PPFD 1000μmol／(m~2·s)左右，气孔导度也未受影响。次高锰处理后，空心莲子草光合速率比对照提高了22．4％，胞间CO_2下降；叶绿素b与总叶绿素含量分别比对照提高了20．7％和17．9％，促进了其天线色素的转换效率与PSⅡ电子传递速率，空心莲子草用于PSⅡ光化学反应的能量较多。高锰处理后光合速率显著降低，比对照下降了11．8％，胞间CO_2也下降。高锰处理下叶绿素含量以及叶绿素a和叶绿素b的比值与对照相当，可能对原初反应伤害不大，但是导致电子传递受阻，叶片实际光化学效率与电子传递速率降低，PSⅡ开放的反应中心比例与参与CO_2固定的电子减少，也可能导致碳同化降低迫使叶片的PQ库还原程度升高，通过热耗散的能量增加。总体上，次高锰处理促进了空心莲子草的光合作用，而高锰处理一定程度上抑制了空心莲子草的光合作用。
(3)膜脂过氧化与抗氧化酶活性测定结果表明，次高锰处理后40d和120 d，空心莲子草体内膜脂系统正常，叶片中膜脂过氧化产物丙二醛(MDA)含量，以及超氧化物歧化酶(SOD)和抗坏血酸过氧化酶(APX)活性与对照无显著差异。高锰处理后40 d，MDA含量高于对照，SOD活性与对照相当，APX活性提高为对照的216．5％；高锰处理后120 d，MDA含量与对照无显著差异，SOD活性比对照提高了69．0％；APX活性下降为对照的59．0％。同时进行草甘膦(68 g ae／ha)茎叶处理后，不同浓度锰处理间可溶性蛋白含量变化无显著差异；对照空心莲子草的SOD和APX活性在草甘膦处理后8 d显著下降，而草甘膦处理后2 d次高锰条件下SOD活性即下降；高锰条件下SOD、APX活性也已下降，且MDA含量升高幅度比对照大。上述结果表明，次高锰条件下，空心莲子草能启动抗氧化酶系统有效清除自由基，保护膜系统；高锰胁迫初期，SOD、APX等抗氧化酶系统无法有效清除高锰浓度的氧化毒害，膜脂过氧化增加，但适应120 d后能有效清除自由基。此外，草甘膦处理后次高锰与对照植株的自由基清除能力相当，但高锰处理空心莲子草自由基清除能力低于对照。
(4)空心莲子草植株不同部位9种元素测定结果表明，空心莲子草根系吸收锰后向地上部运转，过量的锰主要积累在地下组织和叶片中。对照植株培养120 d，叶片、茎与地下组织中锰含量分别为0．13、0．042和0．15 mg／g，而次高锰与高锰处理叶片中锰含量分别为对照的5．2倍和44．6倍，地下组织中分别是对照的5．0倍与45．3倍。对照植株叶片中铁、铜、锌、硼与钼的含量分别为163．0、15．7、74．0、0．7和34．4μg／g，次高锰处理叶片中五种元素的含量显著提高，分别比对照提高了21．2％、17．1％、5．0％、47．9％和33．9％，高锰处理叶片中五种元素的含量分别比对照提高了12．5％、25．3％、7．9％、73．1％和43．4％，也均显著高于对照。对照植株地下组织中钙、镁和钾的含量分别为30．1、3．4、30．1 mg／g，次高锰处理地下组织中仅钾含量比对照降低了25．0％，高锰处理地下组织中三种元素含量分别比对照降低了44．7％、31．1％和19．0％，均为显著差异。上述结果表明，次高锰和高锰条件下促进了空心莲子草对铁、铜、锌、硼与钼的吸收，但高锰处理抑制了空心莲子草对钙、镁和钾的吸收。
(5)高锰胁迫下空心莲子草对草甘膦产生了耐药性，且随着锰浓度提高耐药性增强。次高锰与高锰培养120 d，草甘膦(68 g ae／ha)处理后20 d，茎叶鲜重抑制率分别比常规浓度锰处理(90．9％)下降了17．4与44．6个百分点，草甘膦处理后50 d存活的地下主根茎数分别是对照(25．0％)的3．0和3．6倍。次高锰与高锰条件下，草甘膦对空心莲子草地下根茎的ED_(50)值分别是对照(48．1 g ae／ha)的1．5和2．4倍。同位素示踪结果表明，在空心莲子草顶端往下第四对叶进行~(14)C-草甘膦点叶处理后3 d，次高锰与高锰胁迫下空心莲子草叶片的药剂吸收未减少。次高锰条件下~(14)C-草甘膦被叶片吸收后输导至植株其它部位的量比对照(处理剂量的14．7％)提高了53．1％，~(14)C-草甘膦在点药处理叶以上茎叶与根茎中的浓度分别比对照提高了59．7％与37．5％。高锰条件下~(14)C-草甘膦处理后积累在处理叶中未向外输导的药剂量比对照增加了50．0％，但未减少在地下组织的积累。药剂处理后7 d，放射性成像图显示，次高锰与高锰条件下处理叶以下叶片中~(14)C-草甘膦的积累量比对照少。在~(14)C-草甘膦处理后14 d，采用薄层层析法从地下组织中提取分离到的~(14)C-草甘膦残留量约为每株杂草10 ng左右，占~(14)C-草甘膦处理剂量的0．8％左右，~(14)C-草甘膦代谢产物氨甲基磷酸(AMPA)未检出。上述结果表明，次高锰条件下~(14)C-草甘膦在空心莲子草中的传导量增加，地下根茎中的~(14)C-草甘膦含量提高；高锰条件下空心莲子草根茎与根系中的草甘膦积累未减少；草甘膦在不同锰浓度下培养的空心莲子草体内的代谢速度无显著差异。
(6)改进草甘膦的使用技术是治理耐药性空心莲子草的重要措施之一。草甘膦药液在空心莲子草植株上的沉积试验结果表明，用体积中径(VMD)149．5～233．7μm的雾滴喷雾，草甘膦在空心莲子草叶片的沉积量在VMD 1 57．3μm时较多，随着雾滴体积中径进一步增大，沉积量显著减少，VMD 157．3μm处理的草甘膦沉积量比VMD 233．7μm处理的沉积量提高35．1％。相同草甘膦剂量，用较小雾滴(VMD 157．3μm)与较少施药液量(339 L／hm~2)处理的沉积量比较大雾滴(VMD233．4μm)与较多施药液量(694．5 L／hm~2)处理提高了54．2％。施药液量超过382．5L／hm~2后，草甘膦药液的流失明显增多。草甘膦(800 mg ae／L)药液在空心莲子草叶片的最大稳定持留量约为4．92μg ae／cm~2。上述结果表明，喷施草甘膦时采用VMD 149．5～157．3μm的较小雾滴和约339 L／hm~2的施药液量，草甘膦防治空心莲子草时的药剂有效利用率比较高。
Alligator weed,Alternanthera philoxeroides (Mart.)Griseb.,is one of the mostinvasive alien weeds in China.Hydroponic culture experiments were conducted in thepresent study to investigate the growth and propagation of alligator weed,and theassociation between Mn accumulation and absorption of other ions under Mn stress,and the physiological responses of photosynthesis and antioxidant enzymatic systemsof alligator weeds to Mn stress.The susceptibility of alligator weed to glyphosate andthe behavior of ~(14)C-glyphosate in this weed were also analyzed.Furthermore,thedeposition characteristic of glyphosate in alligator weed leaves was determined.Theresults were summarized below:
The growth of alligator weed was significantly enhaced in high Mn concentrationranging from 0.31～2.45 mM treatment as compared to that of the control (0.0091 mMMn),but inhibited when Mn concentration increased to more than 4.90 mM.Thenumber of new rhizomes developed from the parent rhizomes on 40,120,and 365 daysafter treatment (DAT)of 0.31 mM Mn were 2.3～2.6 times of those of the control,andthe biomass of underground parts were significantly increased by 24.1%～32.2%.Theshoot fresh weight per year was significantly increased by 18.4 % as compared to thatof the control.The propagation of alligator weed was also stimulated in 2.45 mM Mntreatment.The number of new rhizomes developed from the parent rhizomes on 40,120,and 365 DAT were 2.4～3.0 times more than that of the control,and the biomassof underground parts increased from 82.2% of the control at 40 DAT to 125.1% of thecontrol at 365 DAT.However,the biomass of shoots per year was only 71.8 % of thecontrol.The results indicate that the potential of alligator weed fitness to Mn stressprobably contribute to its invasive ability.
The light saturation point of alligator weed was not altered under Mn stress,PPFDapproximately around 1000μmolm~(-2)s~(-1).Compared to the control,the content of bothChlorophyll b and total Chlorophyll increased in 0.31 mM Mn treatment,and theexcitation energy capture efficiency of PSⅡreaction centers (F_v′/F_m′)and electrontransport rate (ETR)was improved,with more excited energy used for photochemistry reaction,and the photosynthetic rate (P_n)was also increased by 22.4%.In 2.45 mM Mntreatment,the content of Chlorophyll b and the ratio of Chlorophyll a/b were thesame as the control,but the effective PSII quantum yield (Φ_(PSⅡ))and ETR as well asphotochemical quenching (q_p)declined compared to the control,and consequently CO_2assimilation probably be inhibited.Therefore,the photosynthesis of alligator weed wasimproved under 0.31 mM Mn treatment,but was inhibited under 2.45 mM Mntreatment.
The antioxidant enzymatic systems under high Mn stress were characterized.Thecontent of malondialdehyde (MDA),and the activity of superoxide dismutase (SOD)and ascorbate peroxidase (APX)in alligator weed leaves were similar to those of thecontrol in 0.31 mM Mn treatment.The content of MDA was similar to that of thecontrol,the activity of SOD was significantly increased by 69.0%,and the activity ofAPX was increased and then followed by decrease in 2.45 mM Mn treatment ascompared to those of the control.After glyphosate (68 g ae/ha)treatment,the solubleprotein content was not affected under Mn stress;however,the activity of SODdecreased earlier than that of the control in 0.31 mM Mn treatment;and the MDAcontent increased earlier and the activity of SOD and APX decreased earlier comparedto those of the control in 2.45 mM Mn treatment.The above results indicate that theantioxidant enzymatic systems might have been initiated in alligator weeds in responseto Mn stress.
Mn is absorbed by the roots of alligator weed and then translocated in the plant.Excess Mn is mainly accumulated in leaves and underground parts of the weed.On120 days after 0.0091 mM Mn treatment (the control),the concentration of Mn inleaves,stems and underground parts were 0.13,0.042 and 0.15 mg/g,respectively.In0.31 and 2.45 mM Mn treatments,the concentrations of Mn in leaves were 5.2 and44.6 times those of the control,and the concentrations of Mn in underground partswere 5.0 and 45.3 times those of the control.The concentrations of Iron (Fe),copper(Cu),zinc (Zn),boron (B),and molybdenum (Mo)in leaves of the control were 163.0, 15.7,74.0,0.7,and 34.4μg/g,respectively,however,the concentrations of Fe,Cu,Zn,B,and Mo in leaves were significantly increased by 21.2%,17.1%,5.0%,47.9% and33.9%,respectively,in 0.31 mM Mn treatment and by 12.5%,25.3%,7.9%,73.1%and 43.4%,respectively,in 2.45 mM Mn treatment.The concentration of calcium (Ca),magnesium (Mg)and potassium (K)in underground parts of the control were 30.1,3.4and 30.1 mg/g,respectively.However,the concentration of K in underground partswas decreased by 25.0% in 0.31 mM Mn treatment,and those of Ca,Mg and K inunderground parts were decreased by 44.7%,31.1% and 19.0%,respectively,in 2.45mM Mn treatment.
Alligator weeds showed remarkably variable susceptibility to glyphosate under Mnstress.On 120 days after treatment of 0.31 and 2.45 mM Mn,shoots fresh weightreduction declined significantly by 17.4 % and 44.6 %,respectively,compared to thatof the control on 20 days after glyphosate application (DAA).The number of survivingrhizomes in both high Mn treatments was approximately 3 times more than that of thecontrol at 50 DAA.The ~(14)C-glyphosat absorption by alligator weed was not reducedunder Mn stress 3 days after ~(14)C-glyphosate application,but more ~(14)C-glyphosate wastranslocated.Consequently the ~(14)C-glyphosate concentration in both shoots andrhizomes were improved in 0.31 mM Mn treatment.Though more ~(14)C-glyphosate wasremained in the treated leaves in 2.45 mM Mn treatment,the ~(14)C-glyphosateconcentration in rhizomes and roots were remained at the same level as the control.Less ~(14)C-glyphosate was accumulated in the treated basal leaves under Mn stress 7days after ~(14)C-glyphosate application.About 10 ng per plant of ~(14)C-glyphosate residuewas detected in rhizomes and roots 14 days after ~(14)C-glyphosate application,whichwas approximately 0.8% of total ~(14)C-glyphosate applied.The main metabolic productof ~(14)C-glyphosate——AMPA was not detected using thin-layer chromatography (TLC).These results suggest that the tolerance of alligator weed to glyphosate under Mn stressis probably not due to the decrease of absorption and translocation of glyphosate andmight not caused by rapid metabolism of glyphosate in the weed.
A noteworthy strategy for tolerant or resistant weed management is to improveherbicide application technique.Amogy the droplet sizes ranging from VMD 149.5μmto 233.7μm,the deposition of glyphosate on alligator weed leaves treated in VMD157.3μm was more than that in larger droplets.The deposition of glyphosate treated inless spray volume (339 L/hm~2)and smaller droplets (VMD 157.3μm)was 1.54 timesmore than that in more spray volume (694.5 L/hm~2)and coarse droplets (VMD 233.7μm).The deposition of glyphosate was increased slightly when spray volume wasmore than 382.5 L/hm~2,and the maximum deposition of glyphosate on alligator weedleaves was approximately 4.92μg ae/cm~2 when glyphosate was applied inconcentration of 800 mg ae/L and VMD 149.5μm.The above results indicate that inrelatively smaller droplets of VMD (149.5～157.3μm)and lower spray volume(approximately 339 L/hm~2),the efficiency of glyphosate application would beimproved significantly in the process of alligator weed management.
(1)温室条件下空心莲子草生物量调查结果表明，锰浓度在0．31～2．45 mM范围内，培养100 d后空心莲子草茎叶生物量显著高于常规浓度锰(0．0091 mM，对照)培养的植株，锰浓度超过4．90 mM后，空心莲子草生物量随锰浓度进一步提高而下降。次高锰(0．31 mM)条件下培养40、120、365 d后从扦插主根茎上长出的新根茎数是对照的2．3～2．6倍；地下组织生物量比对照提高了24．1％～32．2％；一年的茎叶生物量之和比对照(17．5 g／株)提高了18．4％，均存在显著差异。高锰(2．45 mM)条件下培养40、120、365 d后新根茎数是对照的2．4～3．0倍；培养40 d时地下组织生物量只有对照的82．2％，尔后逐步增加，培养365 d时比对照增加了25．1％；但一年的茎叶生物量之和显著下降，只有对照的71．8％。本实验结果表明，空心莲子草的入侵能力与土壤中有效锰含量密切相关，次高锰促进了空心莲子草生长和地下根茎形成；高锰有利于地下根茎形成，但抑制了地上部生长。
(2)光合速率与荧光参数测定结果表明，不同浓度锰处理后120 d，空心莲子草的光饱和点不受影响，均在PPFD 1000μmol／(m~2·s)左右，气孔导度也未受影响。次高锰处理后，空心莲子草光合速率比对照提高了22．4％，胞间CO_2下降；叶绿素b与总叶绿素含量分别比对照提高了20．7％和17．9％，促进了其天线色素的转换效率与PSⅡ电子传递速率，空心莲子草用于PSⅡ光化学反应的能量较多。高锰处理后光合速率显著降低，比对照下降了11．8％，胞间CO_2也下降。高锰处理下叶绿素含量以及叶绿素a和叶绿素b的比值与对照相当，可能对原初反应伤害不大，但是导致电子传递受阻，叶片实际光化学效率与电子传递速率降低，PSⅡ开放的反应中心比例与参与CO_2固定的电子减少，也可能导致碳同化降低迫使叶片的PQ库还原程度升高，通过热耗散的能量增加。总体上，次高锰处理促进了空心莲子草的光合作用，而高锰处理一定程度上抑制了空心莲子草的光合作用。
(3)膜脂过氧化与抗氧化酶活性测定结果表明，次高锰处理后40d和120 d，空心莲子草体内膜脂系统正常，叶片中膜脂过氧化产物丙二醛(MDA)含量，以及超氧化物歧化酶(SOD)和抗坏血酸过氧化酶(APX)活性与对照无显著差异。高锰处理后40 d，MDA含量高于对照，SOD活性与对照相当，APX活性提高为对照的216．5％；高锰处理后120 d，MDA含量与对照无显著差异，SOD活性比对照提高了69．0％；APX活性下降为对照的59．0％。同时进行草甘膦(68 g ae／ha)茎叶处理后，不同浓度锰处理间可溶性蛋白含量变化无显著差异；对照空心莲子草的SOD和APX活性在草甘膦处理后8 d显著下降，而草甘膦处理后2 d次高锰条件下SOD活性即下降；高锰条件下SOD、APX活性也已下降，且MDA含量升高幅度比对照大。上述结果表明，次高锰条件下，空心莲子草能启动抗氧化酶系统有效清除自由基，保护膜系统；高锰胁迫初期，SOD、APX等抗氧化酶系统无法有效清除高锰浓度的氧化毒害，膜脂过氧化增加，但适应120 d后能有效清除自由基。此外，草甘膦处理后次高锰与对照植株的自由基清除能力相当，但高锰处理空心莲子草自由基清除能力低于对照。
(4)空心莲子草植株不同部位9种元素测定结果表明，空心莲子草根系吸收锰后向地上部运转，过量的锰主要积累在地下组织和叶片中。对照植株培养120 d，叶片、茎与地下组织中锰含量分别为0．13、0．042和0．15 mg／g，而次高锰与高锰处理叶片中锰含量分别为对照的5．2倍和44．6倍，地下组织中分别是对照的5．0倍与45．3倍。对照植株叶片中铁、铜、锌、硼与钼的含量分别为163．0、15．7、74．0、0．7和34．4μg／g，次高锰处理叶片中五种元素的含量显著提高，分别比对照提高了21．2％、17．1％、5．0％、47．9％和33．9％，高锰处理叶片中五种元素的含量分别比对照提高了12．5％、25．3％、7．9％、73．1％和43．4％，也均显著高于对照。对照植株地下组织中钙、镁和钾的含量分别为30．1、3．4、30．1 mg／g，次高锰处理地下组织中仅钾含量比对照降低了25．0％，高锰处理地下组织中三种元素含量分别比对照降低了44．7％、31．1％和19．0％，均为显著差异。上述结果表明，次高锰和高锰条件下促进了空心莲子草对铁、铜、锌、硼与钼的吸收，但高锰处理抑制了空心莲子草对钙、镁和钾的吸收。
(5)高锰胁迫下空心莲子草对草甘膦产生了耐药性，且随着锰浓度提高耐药性增强。次高锰与高锰培养120 d，草甘膦(68 g ae／ha)处理后20 d，茎叶鲜重抑制率分别比常规浓度锰处理(90．9％)下降了17．4与44．6个百分点，草甘膦处理后50 d存活的地下主根茎数分别是对照(25．0％)的3．0和3．6倍。次高锰与高锰条件下，草甘膦对空心莲子草地下根茎的ED_(50)值分别是对照(48．1 g ae／ha)的1．5和2．4倍。同位素示踪结果表明，在空心莲子草顶端往下第四对叶进行~(14)C-草甘膦点叶处理后3 d，次高锰与高锰胁迫下空心莲子草叶片的药剂吸收未减少。次高锰条件下~(14)C-草甘膦被叶片吸收后输导至植株其它部位的量比对照(处理剂量的14．7％)提高了53．1％，~(14)C-草甘膦在点药处理叶以上茎叶与根茎中的浓度分别比对照提高了59．7％与37．5％。高锰条件下~(14)C-草甘膦处理后积累在处理叶中未向外输导的药剂量比对照增加了50．0％，但未减少在地下组织的积累。药剂处理后7 d，放射性成像图显示，次高锰与高锰条件下处理叶以下叶片中~(14)C-草甘膦的积累量比对照少。在~(14)C-草甘膦处理后14 d，采用薄层层析法从地下组织中提取分离到的~(14)C-草甘膦残留量约为每株杂草10 ng左右，占~(14)C-草甘膦处理剂量的0．8％左右，~(14)C-草甘膦代谢产物氨甲基磷酸(AMPA)未检出。上述结果表明，次高锰条件下~(14)C-草甘膦在空心莲子草中的传导量增加，地下根茎中的~(14)C-草甘膦含量提高；高锰条件下空心莲子草根茎与根系中的草甘膦积累未减少；草甘膦在不同锰浓度下培养的空心莲子草体内的代谢速度无显著差异。
(6)改进草甘膦的使用技术是治理耐药性空心莲子草的重要措施之一。草甘膦药液在空心莲子草植株上的沉积试验结果表明，用体积中径(VMD)149．5～233．7μm的雾滴喷雾，草甘膦在空心莲子草叶片的沉积量在VMD 1 57．3μm时较多，随着雾滴体积中径进一步增大，沉积量显著减少，VMD 157．3μm处理的草甘膦沉积量比VMD 233．7μm处理的沉积量提高35．1％。相同草甘膦剂量，用较小雾滴(VMD 157．3μm)与较少施药液量(339 L／hm~2)处理的沉积量比较大雾滴(VMD233．4μm)与较多施药液量(694．5 L／hm~2)处理提高了54．2％。施药液量超过382．5L／hm~2后，草甘膦药液的流失明显增多。草甘膦(800 mg ae／L)药液在空心莲子草叶片的最大稳定持留量约为4．92μg ae／cm~2。上述结果表明，喷施草甘膦时采用VMD 149．5～157．3μm的较小雾滴和约339 L／hm~2的施药液量，草甘膦防治空心莲子草时的药剂有效利用率比较高。
Alligator weed,Alternanthera philoxeroides (Mart.)Griseb.,is one of the mostinvasive alien weeds in China.Hydroponic culture experiments were conducted in thepresent study to investigate the growth and propagation of alligator weed,and theassociation between Mn accumulation and absorption of other ions under Mn stress,and the physiological responses of photosynthesis and antioxidant enzymatic systemsof alligator weeds to Mn stress.The susceptibility of alligator weed to glyphosate andthe behavior of ~(14)C-glyphosate in this weed were also analyzed.Furthermore,thedeposition characteristic of glyphosate in alligator weed leaves was determined.Theresults were summarized below:
The growth of alligator weed was significantly enhaced in high Mn concentrationranging from 0.31～2.45 mM treatment as compared to that of the control (0.0091 mMMn),but inhibited when Mn concentration increased to more than 4.90 mM.Thenumber of new rhizomes developed from the parent rhizomes on 40,120,and 365 daysafter treatment (DAT)of 0.31 mM Mn were 2.3～2.6 times of those of the control,andthe biomass of underground parts were significantly increased by 24.1%～32.2%.Theshoot fresh weight per year was significantly increased by 18.4 % as compared to thatof the control.The propagation of alligator weed was also stimulated in 2.45 mM Mntreatment.The number of new rhizomes developed from the parent rhizomes on 40,120,and 365 DAT were 2.4～3.0 times more than that of the control,and the biomassof underground parts increased from 82.2% of the control at 40 DAT to 125.1% of thecontrol at 365 DAT.However,the biomass of shoots per year was only 71.8 % of thecontrol.The results indicate that the potential of alligator weed fitness to Mn stressprobably contribute to its invasive ability.
The light saturation point of alligator weed was not altered under Mn stress,PPFDapproximately around 1000μmolm~(-2)s~(-1).Compared to the control,the content of bothChlorophyll b and total Chlorophyll increased in 0.31 mM Mn treatment,and theexcitation energy capture efficiency of PSⅡreaction centers (F_v′/F_m′)and electrontransport rate (ETR)was improved,with more excited energy used for photochemistry reaction,and the photosynthetic rate (P_n)was also increased by 22.4%.In 2.45 mM Mntreatment,the content of Chlorophyll b and the ratio of Chlorophyll a/b were thesame as the control,but the effective PSII quantum yield (Φ_(PSⅡ))and ETR as well asphotochemical quenching (q_p)declined compared to the control,and consequently CO_2assimilation probably be inhibited.Therefore,the photosynthesis of alligator weed wasimproved under 0.31 mM Mn treatment,but was inhibited under 2.45 mM Mntreatment.
The antioxidant enzymatic systems under high Mn stress were characterized.Thecontent of malondialdehyde (MDA),and the activity of superoxide dismutase (SOD)and ascorbate peroxidase (APX)in alligator weed leaves were similar to those of thecontrol in 0.31 mM Mn treatment.The content of MDA was similar to that of thecontrol,the activity of SOD was significantly increased by 69.0%,and the activity ofAPX was increased and then followed by decrease in 2.45 mM Mn treatment ascompared to those of the control.After glyphosate (68 g ae/ha)treatment,the solubleprotein content was not affected under Mn stress;however,the activity of SODdecreased earlier than that of the control in 0.31 mM Mn treatment;and the MDAcontent increased earlier and the activity of SOD and APX decreased earlier comparedto those of the control in 2.45 mM Mn treatment.The above results indicate that theantioxidant enzymatic systems might have been initiated in alligator weeds in responseto Mn stress.
Mn is absorbed by the roots of alligator weed and then translocated in the plant.Excess Mn is mainly accumulated in leaves and underground parts of the weed.On120 days after 0.0091 mM Mn treatment (the control),the concentration of Mn inleaves,stems and underground parts were 0.13,0.042 and 0.15 mg/g,respectively.In0.31 and 2.45 mM Mn treatments,the concentrations of Mn in leaves were 5.2 and44.6 times those of the control,and the concentrations of Mn in underground partswere 5.0 and 45.3 times those of the control.The concentrations of Iron (Fe),copper(Cu),zinc (Zn),boron (B),and molybdenum (Mo)in leaves of the control were 163.0, 15.7,74.0,0.7,and 34.4μg/g,respectively,however,the concentrations of Fe,Cu,Zn,B,and Mo in leaves were significantly increased by 21.2%,17.1%,5.0%,47.9% and33.9%,respectively,in 0.31 mM Mn treatment and by 12.5%,25.3%,7.9%,73.1%and 43.4%,respectively,in 2.45 mM Mn treatment.The concentration of calcium (Ca),magnesium (Mg)and potassium (K)in underground parts of the control were 30.1,3.4and 30.1 mg/g,respectively.However,the concentration of K in underground partswas decreased by 25.0% in 0.31 mM Mn treatment,and those of Ca,Mg and K inunderground parts were decreased by 44.7%,31.1% and 19.0%,respectively,in 2.45mM Mn treatment.
Alligator weeds showed remarkably variable susceptibility to glyphosate under Mnstress.On 120 days after treatment of 0.31 and 2.45 mM Mn,shoots fresh weightreduction declined significantly by 17.4 % and 44.6 %,respectively,compared to thatof the control on 20 days after glyphosate application (DAA).The number of survivingrhizomes in both high Mn treatments was approximately 3 times more than that of thecontrol at 50 DAA.The ~(14)C-glyphosat absorption by alligator weed was not reducedunder Mn stress 3 days after ~(14)C-glyphosate application,but more ~(14)C-glyphosate wastranslocated.Consequently the ~(14)C-glyphosate concentration in both shoots andrhizomes were improved in 0.31 mM Mn treatment.Though more ~(14)C-glyphosate wasremained in the treated leaves in 2.45 mM Mn treatment,the ~(14)C-glyphosateconcentration in rhizomes and roots were remained at the same level as the control.Less ~(14)C-glyphosate was accumulated in the treated basal leaves under Mn stress 7days after ~(14)C-glyphosate application.About 10 ng per plant of ~(14)C-glyphosate residuewas detected in rhizomes and roots 14 days after ~(14)C-glyphosate application,whichwas approximately 0.8% of total ~(14)C-glyphosate applied.The main metabolic productof ~(14)C-glyphosate——AMPA was not detected using thin-layer chromatography (TLC).These results suggest that the tolerance of alligator weed to glyphosate under Mn stressis probably not due to the decrease of absorption and translocation of glyphosate andmight not caused by rapid metabolism of glyphosate in the weed.
A noteworthy strategy for tolerant or resistant weed management is to improveherbicide application technique.Amogy the droplet sizes ranging from VMD 149.5μmto 233.7μm,the deposition of glyphosate on alligator weed leaves treated in VMD157.3μm was more than that in larger droplets.The deposition of glyphosate treated inless spray volume (339 L/hm~2)and smaller droplets (VMD 157.3μm)was 1.54 timesmore than that in more spray volume (694.5 L/hm~2)and coarse droplets (VMD 233.7μm).The deposition of glyphosate was increased slightly when spray volume wasmore than 382.5 L/hm~2,and the maximum deposition of glyphosate on alligator weedleaves was approximately 4.92μg ae/cm~2 when glyphosate was applied inconcentration of 800 mg ae/L and VMD 149.5μm.The above results indicate that inrelatively smaller droplets of VMD (149.5～157.3μm)and lower spray volume(approximately 339 L/hm~2),the efficiency of glyphosate application would beimproved significantly in the process of alligator weed management.
引文
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