乙羧氟草醚和乙草胺对葡萄的危害原因探究
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摘要
葡萄园杂草控制是土壤管理的一个重要方面。近年来由于劳动力紧缺以及缺乏对果园生草的认知，除草剂的使用日趋普及和常规。正因为如此，有关除草剂药害的报道也越来越多。2009年调查中发现山东省曲阜市常年喷施高浓度复合除草剂的巨峰葡萄园中叶片变黑，植株生长势较弱，以此为背景，本文通过大田和盆栽两种方式验证了除草剂对葡萄的伤害，并探究了乙羧氟草醚和乙草胺这两种单剂对葡萄伤害的生理机制以及导致葡萄叶片变黑的原因。主要研究结果如下：1．复合除草剂对田间巨峰葡萄叶片光合作用伤害的验证
在山东省曲阜市，与人工除草的巨峰葡萄园相比，2009年使用除草剂（百草枯、乙草胺和乙羧氟草醚复合）的葡萄园叶片净光合速率（Pn）明显降低，光系统II（PSII）最大光化学效率（Fv/Fm）和性能指数PIABS降低，快速叶绿素荧光诱导动力学曲线中J点和K点荧光上升，表明叶片PSII反应中心和放氧复合体（OEC）受到了损伤。此外，使用除草剂的葡萄园，落叶后枝条可溶性糖、淀粉、游离氨基酸和可溶性蛋白含量较低。
2010年选择使用除草剂但未发生药害的巨峰葡萄园，证实喷施低浓度除草剂（百草枯+乙草胺+乙羧氟草醚复合）降低了前期葡萄中部叶片Pn；而使用复合高浓度除草剂不仅降低了整个生长季中部叶片Pn，而且降低了葡萄果实可溶性固形物和花青素含量以及落叶后枝条中游离氨基酸和可溶性蛋白含量。2．乙草胺和乙羧氟草醚单剂对葡萄叶片光合作用的影响
基质盆栽巨峰葡萄中部叶片Pn随着处理时间延长呈下降趋势，且随着乙草胺和乙羧氟草醚单剂浓度的升高而降低。第30天时，不同部位叶片Pn也均随除草剂浓度的升高而降低，大田推荐最大用量乙草胺处理的上、下部叶Pn显著低于清水对照，而大田推荐最大用量乙羧氟草醚处理的仅中部叶Pn显著降低；高于大田推荐最大用量除草剂处理不同部位叶片Pn均降低，当浓度为田间推荐最大用量10倍时，Pn降低幅度最大。
沙培巨峰叶片光合功能对乙草胺和乙羧氟草醚单剂（浓度均为田间最大推荐用量5倍）均有明显的时空响应效应。喷施初期（处理后第13天），上部叶片对乙草胺反应明显，Pn、Gs、Fv/Fm和实际光化学效率（ΦPSII）显著下降；J点和K点荧光显著上升，PIABS、反应中心吸收的光能用于电子传递的量子产额（φEo）和反应中心捕获的激子中用来推动电子传递的效率（φo）显著下降，这表明PSII反应中心和OEC受损伤，电子传递能力降低，且程度显著高于中部叶片，但随着处理时间的延长，受损伤的程度减轻；在后期（处理后第60天），上部叶与中部叶各指标之间的差距变小；下部叶片对除草剂的响应时间滞后，后期时PSII反应中心和OEC受损伤程度以及电子传递受抑制程度高于中上部叶片。乙羧氟草醚处理对叶片光合作用的影响小于乙草胺处理。
3．乙羧氟草醚和乙草胺单剂对巨峰葡萄叶片颜色的影响
乙羧氟草醚（浓度为田间最大推荐用量5倍）处理后第60天时，沙培巨峰葡萄中、上部叶片变黑，色素含量显著增加，其中上部叶色素含量增加与钙和铁含量增加以及叶绿体数目增多呈显著正相关。而乙草胺（浓度也为田间最大推荐用量5倍）处理后第60天时，中、上部叶片明显变黄，色素含量降低，其中上部叶色素含量降低与锌含量降低呈显著正相关。
4．乙羧氟草醚和乙草胺单剂对巨峰葡萄叶片细胞超微结构的影响
（1）乙羧氟草醚（浓度为田间最大推荐用量5倍）处理后第13天时，沙培巨峰新叶出现褐色斑点，电镜观察结果表明其细胞内膜结构有溶解现象，有一些叶绿体内部基粒和片层结构清晰，还有一些内部模糊；叶绿体、淀粉粒显著减少，且淀粉粒较小。第60天时，细胞膜较完整，而线粒体膜、核膜都较模糊。中部叶大部分叶绿体片层垛叠为基粒，清晰可见，而有少数叶绿体片层间隙很大，叶绿体、淀粉粒显著增多且前者明显变大；上部叶片的叶绿体内部片层间隙较大，叶绿体增多且长度显著增加，但淀粉粒仍较少。
透射电镜观察2011年超高浓度乙羧氟草醚处理巨峰葡萄重新萌芽后的畸形叶片，发现深绿色部位细胞膜结构完整，与同等大小对照叶片相比，叶绿体、淀粉粒和基粒显著增多，且淀粉粒明显变大，而片层数较少，但这些指标除叶绿体长宽外其余均显著高于黄色部位；黄色部位有一些叶绿体片层较清晰，但片层间隙明显变大，还有一些叶绿体内部有溶解的现象，叶绿体和片层与对照相比显著减少，而淀粉粒和基粒显著增多，叶绿体增大变圆，淀粉粒也增大。
（2）乙草胺（浓度为田间最大推荐用量5倍）处理后第60天时，沙培巨峰中部叶细胞内膜结构受损，叶绿体内片层间隙变大，淀粉粒显著增多；上部叶叶绿体内部模糊，且叶绿体膜开口。中部叶叶绿体宽和上部叶叶绿体长分别比对照小21.4%和16.2%。
5．乙草胺和乙羧氟草醚单剂对葡萄叶片抗氧化系统的影响
（1）超高浓度的乙草胺和乙羧氟草醚处理加快了土培巨峰叶片超氧阴离子产生速率，导致膜脂过氧化程度加剧，产物丙二醛增加，这可能与其较低的超氧化物歧化酶、过氧化氢酶、过氧化物酶和抗坏血酸过氧化物酶活性有关。
（2）高浓度乙羧氟草醚处理后，谷胱甘肽还原酶、抗坏血酸氧化酶活性以及上/中部叶片的谷胱甘肽转移酶、单脱氢抗坏血酸还原酶（MDHAR）活性升高，而下部叶MDHAR和上部叶脱氢抗坏血酸还原酶（DHAR）活性仅在14天时升高，中/下部叶DHAR活性在14天和80天时显著升高。这些酶活性的变化导致叶片中抗坏血酸和谷胱甘肽含量发生相应的变化。
6．乙羧氟草醚和乙草胺单剂对葡萄生长的影响
低于除草推荐浓度的乙羧氟草醚（9.4g ai ha-1）和乙草胺（1123g ai ha-1）显著促进霞多丽/SO4的新梢生长；而高于除草推荐最大用量的乙羧氟草醚（187g ai ha-1）和乙草胺（11230g ai ha-1）降低了根系活力，抑制了新梢的生长，第50天时新梢长度分别比对照低86.6%、70.6%，前者新梢中IAA含量显著降低，而ABA含量显著升高。
高浓度的乙羧氟草醚（187g ai ha-1）和乙草胺（11230g ai ha-1）降低了沙培巨峰葡萄植株总生物量，但仅前者与对照差异显著；乙羧氟草醚处理后，根重显著低于对照，植株根冠比降低；而乙草胺处理后，植株根冠比升高。
7．除草剂的后效
在使用较高浓度复合除草剂的葡萄园改为人工除草的情况下，巨峰葡萄上、中、下部叶片的Pn与相邻长期人工除草的葡萄园相比降低幅度仍然高达34.9%、28.5%和61.2%，说明除草剂有较长时期的残效作用；盆栽试验也证明，除草推荐上限浓度或更高浓度的乙羧氟草醚处理后，巨峰植株翌年生长受到明显抑制，特别是超高浓度处理的植株，翌年发芽后叶片畸形且不能恢复正常。因此建议严格控制除草剂的使用浓度，有条件的地方采用葡萄园生草。
Weed control is one important aspect of soil managements in vineyard. In recent years,due to labour shortage and lack of knowledge on grass cover in orchards, herbicidesapplication becomes increasingly universal and general in vineyards. Just because of that,there are more and more reports about herbicide phytotoxicity. In the survey of Qufu city,Shandong province in2009, the grape leaves in vineyard became dark green, and plantgrowth power was weak. Upon this background, our study verified the damage of herbicidesto grapevines through field and pot experiments, and explored the damage physiologicalmechanisms of acetochlor and fluoroglycofen to grapevines, so the reasons of grape leavesbecoming dark green. The results were as follows:
1. Verification the combined herbicide reduced photosynthesis in grape leaves of vineyard
In Qufu city, Shandong province, compared with the artificial weeding, the leaf netphotosynthesis rate (Pn) in vineyards applied with combined herbicide (paraquat, acetochlorand fluoroglycofen) in2009decreased significantly; the maximum photochemical efficiencyof PSII (Fv/Fm) and performance index (PIABS) also decreased; however, the relative variablefluorescence in the J step and K step increased, which indicated the PSII reaction center andoxygen-evolving complex (OEC) were damaged. Besides, the soluble sugar, starch, freeamino acids and soluble protein content in branches were lower.
In2010, Kyoho (Vitis vinifera×V. labrusca) vineyard which used herbicides but did nothappen phytotoxicity was selected. Lower concentration of combined herbicides (paraquat,acetochlor and fluoroglycofen) reduced Pn of middle-node leaves in the earlier treatment time;while the higher level reduced Pn through the growing season, so the soluble solid andanthocyanin content of grapes, the free amino acid and soluble protein content of branchesafter defoliation.
2. Effects of acetochlor and fluoroglycofen on photosynthesis in grape leaves
In the soil pot experiments, the Pn in middle-node leaves of Kyoho decreased along withthe processing time, besides, acetochlor and fluoroglycofen reduced Pn in a dose-dependentmanner. In the30thday, Pn in different nodes of leaves also decreased along with acetochlorand fluoroglycofen concentration increased. Under the field maximum recommended dose,acetochlor reduced Pn in upper-and middle-node leaves, while fluoroglycofen only reduced itin middle-node leaves; higher concentration of herbicides significantly reduced Pn in all nodeleaves, which caused the greatest reduction under ten times of field maximum recommendeddose.
The photosynthetic function in Kyoho leaves grown in sand shown obvious time-andspace-effect responding to acetochlor and fluoroglycofen (the concentrations were both fivetimes of field maximum recommended dose). In the earlier treatment time (13thday aftertreatment), the upper-node leaves showed significant responses to acetochlor, the Pn and Gsdecreased significantly, so the Fv/Fm and quantum yield of PSII electron transport; therelative variable fluorescence in the J step and K step of the chlorophyll fluorescencetransients increased obviously, while the PIABS, φEoand φodecreased markedly, whichindicated that the PSII reaction center and OEC were damaged, meanwhile the electrontransport ability decreased; furthermore the damage and reduction were severer than inmiddle-node leaves, but the degree of damage was alleviated along with the treatment time.However, the differences of the photosynthetic and chlorophyll fluorescence parametersbetween upper-and middle-node leaves diminished in the later stage (60thday after treatment).The bottom-node leaves appeared later responses to herbicides than that of theupper-/middle-node leaves; in the later stage, the damaged extent of PSII reaction center andOEC, and the inhibition of electron transport ability were much higher than that ofupper-/middle-node leaves. The effects of fluoroglycofen on photosynthetic apparatus wereless than acetochlor.
3. Effects of fluoroglycofen and acetochlor on colour of Kyoho leaves
In the60thday after fluoroglycofen (the concentrations was five times of field maximumrecommended dose) treatment, the middle-and upper-node leaves became dark green, and thepigments increased obviously. Positive correlations between the increase of Ca, Mg content,chloroplast number and rise of pigment content were observed.
However, in the60thday after acetochlor (the concentrations was also five times of fieldmaximum recommended dose) treatment, the middle-and upper-node leaves of grapevinesgrown in sand became yellow, and the pigments decreased significantly. There was a positivecorrelation between the reduction of Zn content and decrease of pigment contents.
4. Effects of fluoroglycofen and acetochlor on cell ultrastructure of Kyoho leaves
(1) In the13thday after fluoroglycofen (the concentration was five times of fieldmaximum recommended dose) treatment, brown spots appeared in the young leaf ofgrapevines grown in sand. The membrane dissolved, but grana and lamellaes were clear insome chloroplasts with increased gap between lamellaes; while the inside of somechloroplasts became fuzzy. Not only the chloroplast and starch grain number was reducedobviously, but also the starch grains were smaller. In the60thday after treatment, the cellmembrane and chloroplast membrane was integrated, while the mitochondria and nuclearmembrane were impaired. In the middle-node leaves, the grana and lamellae were clear inmost chloroplasts, while the gap between lamellaes in few ones increased; not only thechloroplast and starch grain number per cell increased obviously, but also the size ofchloroplasts enhanced. The chloroplasts of upper-node leaves were same to the control ones,but lamellaes with large gap were observed; the number and length of chloroplasts enhanced;but the starch grain number was still less than that of control.
Transmission electron microscope analysis on the deformed leaves of grapevines treatedwith extremely high level of fluoroglycofen (375g ai ha-1) showed that: a) in dark green parts,membrane was integrated; compared with the same size control leaves, the chloroplast, starchgrain and granum were much more, even the starch grain became obviously larger, however,the lamellae number per granum was44.4%less; but all the datas except chloroplast size werehigher than that in yellow parts; b) in yellow parts, the lamellaes in some chloroplasts wereclear, but the gap between them was obviously larger, while inside of some ones appeareddissolved phenomenon. Compared with the control, the numbers of chloroplast per cell andlamellae per granum reduced; however, the numbers of starch grain per cell and granum perchloroplast increased. Chloroplasts went larger and round, starch grains also became larger.
(2) In the60thday after acetochlor (the concentrations was also five times of fieldmaximum recommended dose) treatment, the cell membrane of middle-node leaves was impaired, the gap between chloroplastic lamellaes and the starch grain number per cellincreased; while in upper-node leaves, the inside of chloroplast became fuzzy with nolamellae, and there was a opening in chloroplastic membrane. The width of chloroplast inmiddle-node leaves and the length of one in upper-node leaves were respectively21.4%,16.2%smaller than that of control.
5. Effects of acetochlor and fluoroglycofen on antioxidant defense system in Kyoho leaves
(1) Higher level of acetochlor and fluoroglycofen accelerated the superoxide productionrate, resulted in aggravation of the membrane lipid peroxidation, led to accumulation of MDA,which might be related with lower antioxidant enzyme (superoxide dismutase, catalase,peroxidase and ascorbic acid peroxidase) activities.
(2) Higher concentration of fluoroglycofen treatment improved the glutathione reductaseand ascorbate oxidase, so the glutathione transferase and monodehydroascorbate reductase(MDHAR) in upper-and middle-node leaves. While the activities of MDHAR in bottom-nodeleaves and dehydroascorbate reductase (DHAR) in upper-node leaves increased only in14thday after treatment, the DHAR activity in middle-and bottom-node leaves increased only inthe14thand80thday. The changes might resulted in relative changes of ascorbic acid andglutathione content.
6. Effects of fluoroglycofen and acetochlor on shoot growth of grapevines
Fluoroglycofen (9.4g ai ha-1) and acetochlor (1123g ai ha-1) significantly improved thegrowth of new shoots in Chardonnay/SO4, while187g ai ha-1fluoroglycofen and11230g aiha-1acetochlor reduced root activity and suppressed the growth of shoots, i.e. by86.6%,70.6%respectively in the50thday. Compared with the control, IAA content in187g ai ha-1fluoroglycofen groups was significantly lower than control, while ABA content wassignificantly higher.
Fluoroglycofen (187.5g ai ha-1) and acetochlor (11230g ai ha-1) reduced the total plantbiomass of Kyoho grapevines grown in sand, but there was significant difference onlybetween the former groups and control. Fluoroglycofen obviously reduced the root weightand root-shoot ratio, while acetochlor caused increase of root-shoot ratio.
7. Aftereffects of herbicides
After the artifical weeding instead of higher concentration of combined herbicide application in2010, compared with the adjacent vineyard which used artificial weeding for along time, the reductions in Pn of upper-, middle-and bottom-node leaves were still high(34.9%,28.5%,61.2%respectively). It is indicated that herbicides have a relatively longperiod effects. And in the pot experiments, application of fluoroglycofen under the dose or theones higher than field maximum recommended in first year obviously inhibited the growth ofKyoho in the second year, especially375g ai ha-1fluoroglycofen caused the leaf deformedand could not return to normal situation. It is suggested that the orchardmen should controlthe dosage of herbicide strictly, and adopt green cover in conditional place.
在山东省曲阜市，与人工除草的巨峰葡萄园相比，2009年使用除草剂（百草枯、乙草胺和乙羧氟草醚复合）的葡萄园叶片净光合速率（Pn）明显降低，光系统II（PSII）最大光化学效率（Fv/Fm）和性能指数PIABS降低，快速叶绿素荧光诱导动力学曲线中J点和K点荧光上升，表明叶片PSII反应中心和放氧复合体（OEC）受到了损伤。此外，使用除草剂的葡萄园，落叶后枝条可溶性糖、淀粉、游离氨基酸和可溶性蛋白含量较低。
2010年选择使用除草剂但未发生药害的巨峰葡萄园，证实喷施低浓度除草剂（百草枯+乙草胺+乙羧氟草醚复合）降低了前期葡萄中部叶片Pn；而使用复合高浓度除草剂不仅降低了整个生长季中部叶片Pn，而且降低了葡萄果实可溶性固形物和花青素含量以及落叶后枝条中游离氨基酸和可溶性蛋白含量。2．乙草胺和乙羧氟草醚单剂对葡萄叶片光合作用的影响
基质盆栽巨峰葡萄中部叶片Pn随着处理时间延长呈下降趋势，且随着乙草胺和乙羧氟草醚单剂浓度的升高而降低。第30天时，不同部位叶片Pn也均随除草剂浓度的升高而降低，大田推荐最大用量乙草胺处理的上、下部叶Pn显著低于清水对照，而大田推荐最大用量乙羧氟草醚处理的仅中部叶Pn显著降低；高于大田推荐最大用量除草剂处理不同部位叶片Pn均降低，当浓度为田间推荐最大用量10倍时，Pn降低幅度最大。
沙培巨峰叶片光合功能对乙草胺和乙羧氟草醚单剂（浓度均为田间最大推荐用量5倍）均有明显的时空响应效应。喷施初期（处理后第13天），上部叶片对乙草胺反应明显，Pn、Gs、Fv/Fm和实际光化学效率（ΦPSII）显著下降；J点和K点荧光显著上升，PIABS、反应中心吸收的光能用于电子传递的量子产额（φEo）和反应中心捕获的激子中用来推动电子传递的效率（φo）显著下降，这表明PSII反应中心和OEC受损伤，电子传递能力降低，且程度显著高于中部叶片，但随着处理时间的延长，受损伤的程度减轻；在后期（处理后第60天），上部叶与中部叶各指标之间的差距变小；下部叶片对除草剂的响应时间滞后，后期时PSII反应中心和OEC受损伤程度以及电子传递受抑制程度高于中上部叶片。乙羧氟草醚处理对叶片光合作用的影响小于乙草胺处理。
3．乙羧氟草醚和乙草胺单剂对巨峰葡萄叶片颜色的影响
乙羧氟草醚（浓度为田间最大推荐用量5倍）处理后第60天时，沙培巨峰葡萄中、上部叶片变黑，色素含量显著增加，其中上部叶色素含量增加与钙和铁含量增加以及叶绿体数目增多呈显著正相关。而乙草胺（浓度也为田间最大推荐用量5倍）处理后第60天时，中、上部叶片明显变黄，色素含量降低，其中上部叶色素含量降低与锌含量降低呈显著正相关。
4．乙羧氟草醚和乙草胺单剂对巨峰葡萄叶片细胞超微结构的影响
（1）乙羧氟草醚（浓度为田间最大推荐用量5倍）处理后第13天时，沙培巨峰新叶出现褐色斑点，电镜观察结果表明其细胞内膜结构有溶解现象，有一些叶绿体内部基粒和片层结构清晰，还有一些内部模糊；叶绿体、淀粉粒显著减少，且淀粉粒较小。第60天时，细胞膜较完整，而线粒体膜、核膜都较模糊。中部叶大部分叶绿体片层垛叠为基粒，清晰可见，而有少数叶绿体片层间隙很大，叶绿体、淀粉粒显著增多且前者明显变大；上部叶片的叶绿体内部片层间隙较大，叶绿体增多且长度显著增加，但淀粉粒仍较少。
透射电镜观察2011年超高浓度乙羧氟草醚处理巨峰葡萄重新萌芽后的畸形叶片，发现深绿色部位细胞膜结构完整，与同等大小对照叶片相比，叶绿体、淀粉粒和基粒显著增多，且淀粉粒明显变大，而片层数较少，但这些指标除叶绿体长宽外其余均显著高于黄色部位；黄色部位有一些叶绿体片层较清晰，但片层间隙明显变大，还有一些叶绿体内部有溶解的现象，叶绿体和片层与对照相比显著减少，而淀粉粒和基粒显著增多，叶绿体增大变圆，淀粉粒也增大。
（2）乙草胺（浓度为田间最大推荐用量5倍）处理后第60天时，沙培巨峰中部叶细胞内膜结构受损，叶绿体内片层间隙变大，淀粉粒显著增多；上部叶叶绿体内部模糊，且叶绿体膜开口。中部叶叶绿体宽和上部叶叶绿体长分别比对照小21.4%和16.2%。
5．乙草胺和乙羧氟草醚单剂对葡萄叶片抗氧化系统的影响
（1）超高浓度的乙草胺和乙羧氟草醚处理加快了土培巨峰叶片超氧阴离子产生速率，导致膜脂过氧化程度加剧，产物丙二醛增加，这可能与其较低的超氧化物歧化酶、过氧化氢酶、过氧化物酶和抗坏血酸过氧化物酶活性有关。
（2）高浓度乙羧氟草醚处理后，谷胱甘肽还原酶、抗坏血酸氧化酶活性以及上/中部叶片的谷胱甘肽转移酶、单脱氢抗坏血酸还原酶（MDHAR）活性升高，而下部叶MDHAR和上部叶脱氢抗坏血酸还原酶（DHAR）活性仅在14天时升高，中/下部叶DHAR活性在14天和80天时显著升高。这些酶活性的变化导致叶片中抗坏血酸和谷胱甘肽含量发生相应的变化。
6．乙羧氟草醚和乙草胺单剂对葡萄生长的影响
低于除草推荐浓度的乙羧氟草醚（9.4g ai ha-1）和乙草胺（1123g ai ha-1）显著促进霞多丽/SO4的新梢生长；而高于除草推荐最大用量的乙羧氟草醚（187g ai ha-1）和乙草胺（11230g ai ha-1）降低了根系活力，抑制了新梢的生长，第50天时新梢长度分别比对照低86.6%、70.6%，前者新梢中IAA含量显著降低，而ABA含量显著升高。
高浓度的乙羧氟草醚（187g ai ha-1）和乙草胺（11230g ai ha-1）降低了沙培巨峰葡萄植株总生物量，但仅前者与对照差异显著；乙羧氟草醚处理后，根重显著低于对照，植株根冠比降低；而乙草胺处理后，植株根冠比升高。
7．除草剂的后效
在使用较高浓度复合除草剂的葡萄园改为人工除草的情况下，巨峰葡萄上、中、下部叶片的Pn与相邻长期人工除草的葡萄园相比降低幅度仍然高达34.9%、28.5%和61.2%，说明除草剂有较长时期的残效作用；盆栽试验也证明，除草推荐上限浓度或更高浓度的乙羧氟草醚处理后，巨峰植株翌年生长受到明显抑制，特别是超高浓度处理的植株，翌年发芽后叶片畸形且不能恢复正常。因此建议严格控制除草剂的使用浓度，有条件的地方采用葡萄园生草。
Weed control is one important aspect of soil managements in vineyard. In recent years,due to labour shortage and lack of knowledge on grass cover in orchards, herbicidesapplication becomes increasingly universal and general in vineyards. Just because of that,there are more and more reports about herbicide phytotoxicity. In the survey of Qufu city,Shandong province in2009, the grape leaves in vineyard became dark green, and plantgrowth power was weak. Upon this background, our study verified the damage of herbicidesto grapevines through field and pot experiments, and explored the damage physiologicalmechanisms of acetochlor and fluoroglycofen to grapevines, so the reasons of grape leavesbecoming dark green. The results were as follows:
1. Verification the combined herbicide reduced photosynthesis in grape leaves of vineyard
In Qufu city, Shandong province, compared with the artificial weeding, the leaf netphotosynthesis rate (Pn) in vineyards applied with combined herbicide (paraquat, acetochlorand fluoroglycofen) in2009decreased significantly; the maximum photochemical efficiencyof PSII (Fv/Fm) and performance index (PIABS) also decreased; however, the relative variablefluorescence in the J step and K step increased, which indicated the PSII reaction center andoxygen-evolving complex (OEC) were damaged. Besides, the soluble sugar, starch, freeamino acids and soluble protein content in branches were lower.
In2010, Kyoho (Vitis vinifera×V. labrusca) vineyard which used herbicides but did nothappen phytotoxicity was selected. Lower concentration of combined herbicides (paraquat,acetochlor and fluoroglycofen) reduced Pn of middle-node leaves in the earlier treatment time;while the higher level reduced Pn through the growing season, so the soluble solid andanthocyanin content of grapes, the free amino acid and soluble protein content of branchesafter defoliation.
2. Effects of acetochlor and fluoroglycofen on photosynthesis in grape leaves
In the soil pot experiments, the Pn in middle-node leaves of Kyoho decreased along withthe processing time, besides, acetochlor and fluoroglycofen reduced Pn in a dose-dependentmanner. In the30thday, Pn in different nodes of leaves also decreased along with acetochlorand fluoroglycofen concentration increased. Under the field maximum recommended dose,acetochlor reduced Pn in upper-and middle-node leaves, while fluoroglycofen only reduced itin middle-node leaves; higher concentration of herbicides significantly reduced Pn in all nodeleaves, which caused the greatest reduction under ten times of field maximum recommendeddose.
The photosynthetic function in Kyoho leaves grown in sand shown obvious time-andspace-effect responding to acetochlor and fluoroglycofen (the concentrations were both fivetimes of field maximum recommended dose). In the earlier treatment time (13thday aftertreatment), the upper-node leaves showed significant responses to acetochlor, the Pn and Gsdecreased significantly, so the Fv/Fm and quantum yield of PSII electron transport; therelative variable fluorescence in the J step and K step of the chlorophyll fluorescencetransients increased obviously, while the PIABS, φEoand φodecreased markedly, whichindicated that the PSII reaction center and OEC were damaged, meanwhile the electrontransport ability decreased; furthermore the damage and reduction were severer than inmiddle-node leaves, but the degree of damage was alleviated along with the treatment time.However, the differences of the photosynthetic and chlorophyll fluorescence parametersbetween upper-and middle-node leaves diminished in the later stage (60thday after treatment).The bottom-node leaves appeared later responses to herbicides than that of theupper-/middle-node leaves; in the later stage, the damaged extent of PSII reaction center andOEC, and the inhibition of electron transport ability were much higher than that ofupper-/middle-node leaves. The effects of fluoroglycofen on photosynthetic apparatus wereless than acetochlor.
3. Effects of fluoroglycofen and acetochlor on colour of Kyoho leaves
In the60thday after fluoroglycofen (the concentrations was five times of field maximumrecommended dose) treatment, the middle-and upper-node leaves became dark green, and thepigments increased obviously. Positive correlations between the increase of Ca, Mg content,chloroplast number and rise of pigment content were observed.
However, in the60thday after acetochlor (the concentrations was also five times of fieldmaximum recommended dose) treatment, the middle-and upper-node leaves of grapevinesgrown in sand became yellow, and the pigments decreased significantly. There was a positivecorrelation between the reduction of Zn content and decrease of pigment contents.
4. Effects of fluoroglycofen and acetochlor on cell ultrastructure of Kyoho leaves
(1) In the13thday after fluoroglycofen (the concentration was five times of fieldmaximum recommended dose) treatment, brown spots appeared in the young leaf ofgrapevines grown in sand. The membrane dissolved, but grana and lamellaes were clear insome chloroplasts with increased gap between lamellaes; while the inside of somechloroplasts became fuzzy. Not only the chloroplast and starch grain number was reducedobviously, but also the starch grains were smaller. In the60thday after treatment, the cellmembrane and chloroplast membrane was integrated, while the mitochondria and nuclearmembrane were impaired. In the middle-node leaves, the grana and lamellae were clear inmost chloroplasts, while the gap between lamellaes in few ones increased; not only thechloroplast and starch grain number per cell increased obviously, but also the size ofchloroplasts enhanced. The chloroplasts of upper-node leaves were same to the control ones,but lamellaes with large gap were observed; the number and length of chloroplasts enhanced;but the starch grain number was still less than that of control.
Transmission electron microscope analysis on the deformed leaves of grapevines treatedwith extremely high level of fluoroglycofen (375g ai ha-1) showed that: a) in dark green parts,membrane was integrated; compared with the same size control leaves, the chloroplast, starchgrain and granum were much more, even the starch grain became obviously larger, however,the lamellae number per granum was44.4%less; but all the datas except chloroplast size werehigher than that in yellow parts; b) in yellow parts, the lamellaes in some chloroplasts wereclear, but the gap between them was obviously larger, while inside of some ones appeareddissolved phenomenon. Compared with the control, the numbers of chloroplast per cell andlamellae per granum reduced; however, the numbers of starch grain per cell and granum perchloroplast increased. Chloroplasts went larger and round, starch grains also became larger.
(2) In the60thday after acetochlor (the concentrations was also five times of fieldmaximum recommended dose) treatment, the cell membrane of middle-node leaves was impaired, the gap between chloroplastic lamellaes and the starch grain number per cellincreased; while in upper-node leaves, the inside of chloroplast became fuzzy with nolamellae, and there was a opening in chloroplastic membrane. The width of chloroplast inmiddle-node leaves and the length of one in upper-node leaves were respectively21.4%,16.2%smaller than that of control.
5. Effects of acetochlor and fluoroglycofen on antioxidant defense system in Kyoho leaves
(1) Higher level of acetochlor and fluoroglycofen accelerated the superoxide productionrate, resulted in aggravation of the membrane lipid peroxidation, led to accumulation of MDA,which might be related with lower antioxidant enzyme (superoxide dismutase, catalase,peroxidase and ascorbic acid peroxidase) activities.
(2) Higher concentration of fluoroglycofen treatment improved the glutathione reductaseand ascorbate oxidase, so the glutathione transferase and monodehydroascorbate reductase(MDHAR) in upper-and middle-node leaves. While the activities of MDHAR in bottom-nodeleaves and dehydroascorbate reductase (DHAR) in upper-node leaves increased only in14thday after treatment, the DHAR activity in middle-and bottom-node leaves increased only inthe14thand80thday. The changes might resulted in relative changes of ascorbic acid andglutathione content.
6. Effects of fluoroglycofen and acetochlor on shoot growth of grapevines
Fluoroglycofen (9.4g ai ha-1) and acetochlor (1123g ai ha-1) significantly improved thegrowth of new shoots in Chardonnay/SO4, while187g ai ha-1fluoroglycofen and11230g aiha-1acetochlor reduced root activity and suppressed the growth of shoots, i.e. by86.6%,70.6%respectively in the50thday. Compared with the control, IAA content in187g ai ha-1fluoroglycofen groups was significantly lower than control, while ABA content wassignificantly higher.
Fluoroglycofen (187.5g ai ha-1) and acetochlor (11230g ai ha-1) reduced the total plantbiomass of Kyoho grapevines grown in sand, but there was significant difference onlybetween the former groups and control. Fluoroglycofen obviously reduced the root weightand root-shoot ratio, while acetochlor caused increase of root-shoot ratio.
7. Aftereffects of herbicides
After the artifical weeding instead of higher concentration of combined herbicide application in2010, compared with the adjacent vineyard which used artificial weeding for along time, the reductions in Pn of upper-, middle-and bottom-node leaves were still high(34.9%,28.5%,61.2%respectively). It is indicated that herbicides have a relatively longperiod effects. And in the pot experiments, application of fluoroglycofen under the dose or theones higher than field maximum recommended in first year obviously inhibited the growth ofKyoho in the second year, especially375g ai ha-1fluoroglycofen caused the leaf deformedand could not return to normal situation. It is suggested that the orchardmen should controlthe dosage of herbicide strictly, and adopt green cover in conditional place.
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