丙酯草醚胁迫下油菜与大麦耐性机制及其生理信息的光谱模型构建
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摘要
丙酯草醚是我国具有自主知识产权的农药新品种，能有效防除油菜田间主要单、双子叶杂草。作为一种新型油菜田除草剂，研究其对油菜田阔叶杂草及禾本科杂草的防除效果，明确其作用机制及其对油菜和后茬作物的安全性问题等对该除草剂的田间推广应用具有重要意义。但国内目前对这些方面的研究还不够系统深入。本文即以甘蓝型油菜品种浙双758和大麦品种黄青裸为主要材料，采用生理生化分析方法和显微技术研究了丙酯草醚的作用机理及其胁迫下两种作物的耐性差异机制。为便于该新除草剂应用时对大田油菜生理信息进行活体、实时的监测，本课题将除草剂胁迫下的油菜生理信息与近红外光谱结合，尝试构建了相关生理参数的光谱模型。所取得的主要创新性结果如下：
1、离体及活体条件下，分别研究了丙酯草醚对单双子叶作物及杂草乙酰乳酸合成酶(ALS)活力的影响。在离体条件下，丙酯草醚对ALS活力几乎无抑制效果；而在活体条件下，叶面喷施丙酯草醚对双子叶作物油菜ALS活力抑制作用最小，对单子叶作物大麦抑制作用最显著，对阔叶杂草牛繁缕和禾本科杂草看麦娘的抑制效果介于前两种作物之间。植株不同部位的ALS活力存在较大差异，越幼嫩的部位酶活力越高，因此在ALS活力分析中顶端新叶是合适的研究材料。100mg／L丙酯草醚处理下牛繁缕、油菜和大麦的不同叶位ALS活力变化趋势存在差异，牛繁缕与油菜下层叶的酶活力随处理时间延长而下降的程度比上层叶剧烈，在大麦中则相反；油菜上层叶酶活力随处理时间的延长而迅速恢复，而牛繁缕和大麦则无法恢复。三种不同类型油菜耐药性表现为，芥菜型油菜耐性最强，而白菜型油菜耐性最弱，甘蓝型油菜介于两者之间。
2、探讨了丙酯草醚处理对发芽期油菜及大麦幼苗的生长发育及根细胞活性、有丝分裂、根叶细胞形态及超微结构的影响。种子发芽期是丙酯草醚处理的敏感时期；对发芽期油菜和大麦较为安全的临界浓度为1mg／L；10mg／L以上浓度处理下油菜和大麦幼苗的生长尤其是根系的伸长和发育及根尖细胞活性等方面受到了显著抑制，且抑制效果随处理浓度的升高和处理时间的延长而增大。100mg／L处理下油菜和大麦根尖细胞有丝分裂指数均下降，有丝分裂产生的细胞数量减少造成分生区和伸长区明显缩短；另一方面处理还抑制成熟区细胞的发育，导致如油菜侧根发育明显滞后等现象。100mg／L丙酯草醚处理对油菜和大麦根细胞膜造成较大的破坏并导致胞内电解质外渗；处理还对根细胞中的线粒体外膜造成了一定程度的破坏，并存在胞内微粒体数量增加的现象(推测该除草剂的代谢活化作用可能在微粒体中进行，而线粒体中可能存在其活化后的作用靶标)；丙酯草醚处理显著影响细胞亚结构的发育，如抑制大麦根细胞壁、液泡、质体及其叶细胞中的叶绿体发育，但加速油菜子叶细胞中叶绿体的衰老(推测丙酯草醚具有两重性——加速成熟组织的衰老和延缓幼嫩组织的发育，故可尝试将其开发为一种新型的生长调节剂)。在生化检测及透射电镜(TEM)观察都未发现100mg／L丙酯草醚处理对油菜及大麦根尖细胞的核膜造成明显影响，压片法观察分裂细胞也未发现由处理引起的染色体异常和细胞分裂畸形现象。
3、研究了营养生长期油菜及大麦在丙酯草醚胁迫下植株生物量积累、抗氧化系统和光合系统的差异表现及生理机制。发芽期油菜对丙酯草醚的耐性并不高于大麦，但5叶期油菜植株对丙酯草醚的耐性显著高于大麦，表现在：100 mg／L处理浓度对油菜各生理参数均无显著影响，高浓度处理对油菜植株造成显著影响但抑制作用随时间的延长而减弱：而5叶期大麦对100 mg／L处理浓度已敏感，且抑制效果随处理浓度的升高和时间的延长而加剧。在丙酯草醚胁迫下，大麦过氧化物酶(POD)和超氧化物歧化酶(SOD)活性先显著升高后迅速下降，而油菜却未表现出明显的变化趋势，可见丙酯草醚对大麦的抗氧化系统造成的影响大于油菜；在光合系统方面，丙酯草醚对油菜净光合速率(Pn)几乎无影响但却导致大麦叶片SPAD值和Pn的显著下降，这可能与大麦光合系统Ⅱ(PSⅡ)的非光化学耗散能力(NPQ)受到较大抑制有关；丙酯草醚对大麦光合系统造成较大破坏最终导致其更显著地影响大麦的生物量积累。
4、分析了苗期和抽薹期分别喷施丙酯草醚对油菜产量和种子品质等的影响。明确了在苗期喷施100mg／L丙酯草醚可促进油菜增产且对种子品质无显著影响，但1000mg／L高浓度处理对产量和品质(尤其是蛋白质积累方面)有明显抑制；而油菜抽薹期则是对丙酯草醚处理的敏感时期，即使是100mg／L田间推荐浓度喷施处理也会造成减产并影响种子品质，且处理对产量的影响比对品质的影响更显著，1000mg／L高浓度处理几乎导致颗粒无收。以上不同时期作物间的敏感性差异一方面与作物本身对丙酯草醚的活化和相关代谢途径、能力等存在差异，另一方面也可能与处理方式(根部添加和叶面喷施处理)以及丙酯草醚以根茎吸收为主的特性等原因有关。因此在生产上，作物和杂草的苗龄控制及田间施用方式等都将对丙酯草醚的除草效果及对作物的安全性等方面造成影响，应引起足够重视。
5、将近红外光谱技术用于丙酯草醚胁迫下油菜不同叶位生理信息的监测。成功地对油菜叶片中ALS活力，可溶性蛋白、非可溶性蛋白、总蛋白含量和总氨基酸含量等生理参数进行了光谱建模，模型的预测相关系数皆高于0．9：通过不同光谱预处理、有效波长选择和建模方法筛选等技术手段构建的改良模型预测相关系数高达0．95～0．99。
本研究取得的创新性结果对丙酯草醚对单、双子叶植物抑制机制差异有了更全面深入的认识，这将为该新型油菜田除草剂对油菜及后茬作物大麦等安全性评价方面提供重要参考。本研究中对丙酯草醚处理下油菜叶片的光谱数据与不同生理参数进行建模的尝试，为近红外光谱技术应用于除草剂等各种胁迫下植物药害等的大田活体无损快速监测奠定了较好基础，具有显著的创新性和实践应用意义。
Propyl 4-(2-(4,6-dimethoxypyrimidin-2-yloxy)benzylamino)benzoate (ZJ0273) is a novel herbicideused in the rapeseed field with the advantage of broad weeding spectrum. It is meaningful to study themode of action and tolerance mechanism of this new herbicide among crops and weeds for its extendedapplication. In this study, rapeseed (Brassica napus L. cv. ZS758) and barley (Hordeum vulgare L. cv.Huangqingluo) were investigated to clarify the tolerance mechanism within monocotyledon anddicotyledon species to ZJ0273 stress with various physiological measurements and microscopic analyses.In addition, several near-infrared spectroscopy (NIRS) models were developed to investigate for rapid andnon-destructive determination of physiological characters in rapeseed leaves under herbicide ZJ0273stress. The main results are as follows:
1. The differential tolerance or susceptibility of four plant species viz. B. napus L. (rapeseed),Hordeum vulgare L. (barley), Malachium aquaticum (L.) Fries (water chickweed) and Alopecurusaequalis Sobol. (equal alopecurus), to various concentrations of ZJ0273, was studied based on the rate ofinhibition of acetolactate synthase (ALS) activity. B. napus was observed as the most tolerant among fourspecies in which ALS activity in vivo was hardly affected by ZJ0273 at 100 mg/L (recommended dose inthe rapeseed field). In contrast, Hordeum vulgare was found to be more susceptible while the two weedspecies exhibited moderate susceptibility as compared to Hordeum vulgare. However, ALS activity in vitrowas hardly inhibited by herbicide ZJ0273. In addition, there was a wide variation noted for in vivo ALSactivity among various parts of plants. The uppermost younger leaf seemed to be the most appropriatesample for evaluation, as it showed the highest level of ALS activity in the plant. Spatial and temporalvariations in ALS activity as affected by 100 mg/L ZJ0273 in water chickweed, rapeseed and barley alsosuggested that the synchronized sampling based on the tissue position is necessary to examine precisely,the difference in tolerance or susceptibility to ALS-inhibiting herbicide. The variant tendency of ALSactivity from different parts of plants within monocotyledons and dicotyledons to ZJ0273 treatmentsshowed differential response which revealed some possible mechanisms related to the altered absorptionand translocation of the herbicide. Furthermore, the tolerance among three species of Brassica suggestedthat B. rapa was the most susceptible and B. juncea was the most tolerant to the new herbicide ZJ0273.
2. Response of seedling growth, root cell viability, mitosis, root and leaf cell structure andultrastructure of rapeseed and barley to herbicide ZJ0273 at germination stage was studied to define the action mechanism of this novel herbicide. The results indicated that both of rapeseed and barley seedlingsare very sensitive to ZJ0273 at germination stage and the application of 1 mg/L (optimum concentration) issafer for seedling growth. 10 mg/L ZJ0273 treatment can significantly inhibit seedling growth (especiallythe root elongation and development) and root viability. Besides, the inhibitive effects become intensifiedwith the increase of dosage and duration. Mitotic index of root-tip cells was declined and cell division wasstopped at metaphase after being treated by 100 mg/L ZJ0273, which shortened the meristematic zone andelongation zone in plant root tips. In addition, the cell development in maturation zone was also inhibited,which deferred the development of lateral root. ZJ0273 significantly inhibited the development of cell wall,vacuole, plastid and chloroplast in barley cells while induced the senescence (aging) of chloroplast inrapeseed cotyledon cells, which suggested that the effect of ZJ0273 on cell development is bifacial and itcould be exploited as a plant growth regulator. Furthermore, 100 mg/L ZJ0273 treatment obviouslyinhibited root cell viability and resulted in disintegration of cell and mitochondria membrane, as well asmore microbodies were observed in treated cells, which suggested that ZJ0273 might be activated withinthe microbodies and acting in mitochondria in plant cells. However, nucleus in root and leaf cells ofrapeseed and barley treated by ZJ0273 showed no significant change in comparison to that of control, asshown by the histochemical and transmission electron microscopy (TEM) demonstrations. Furthermore,chromosome number and shape in dividing cells treated by 100 mg/L ZJ0273 were also normal, whichindicated that this new herbicide hardly induced the tolerant weed mutants and it is helpful to itsapplication and commercialization in fields.
3. The tolerance mechanism of rapeseed and barley to ZJ0273 stress was studied by measurements ofbiomass, antioxidants and photosynthetic characteristics. As above results shows, rapeseed is no moretolerant than barley at germination stage. In contrast, the tolerance of rapeseed seedlings at 5-leaf stage issignificantly higher than barley seedlings to ZJ0273, which showed that rapeseed seedlings were hardlyaffected by 100 mg/L treatment where as the inhibitive effect of ZJ0273 treatments at higherconcentrations on physiological characteristics recovered with the passage of time. However, barley wasmore susceptible which was reflected in lower biomass production even under at the lowest rate of 100mg/L of ZJ0273 and the enhanced inhibitive effect along with the increase of treatment concentrations anddurations. In the beginning, rapid increase and then sudden decline of leaf peroxidase (POD) andsuperoxide dismutase (SOD) activities under herbicide stress, indicated that ZJ0273 affected barleyantioxidant system more badly than rapeseed, as compared to the slight changes in rapeseed. In addition,ZJ0273 treatments hardly affected net photosynthesis (Pn) of rapeseed leaves but significantly inhibitedSPAD and Pn values of barley leaves, which indicated that the photosystemⅡof barley was paralyzed byinhibition effect of herbicide on non-photochemical quenching (NPQ) capability and finally, the biomass was reduced more significantly.
4. Effect of herbicide ZJ0273 on plant agronomic characters, seed yield and quality of oilseed rapewas also compared in this study by treating at seedling stage and bolting stage, respectively. The resultsconfirmed that 100 mg/L ZJ0273 (recommended dose in the rapeseed field) promoted seed yield withslight effect on seed quality but significantly reduced rapeseed yield and affected seed quality whenapplied at seedling stage and bolting stage, respectively. The treatment at highest concentration of 1000mg/L applied at seedling stage mainly affected seed quality but badly reduced seed yield when was appliedat bolting stage, which indicated that rapeseed yield and seed quality are more sensitive to ZJ0273 atbolting stage than seedling stage. These results suggested that crop tolerance revealed some possiblemechanisms related to the altered absorption, translocation of the herbicide. In addition, treatmentapproach such as root application or leaf-spray was also possible reason that affected the herbicideinhibitive action according to its root-absorption trait. Thus, both of optimal leaf stage and treatmentapproach should be considered in field application of ZJ0273.
5. Near-infrared spectroscopy (NIRS) was investigated for rapid and non-destructive determination ofvarious physiological characters in rapeseed leaves from different leaf-positions under herbicide stress.NIRS models of ALS activity, soluble, non-soluble, total protein and total amino acid contents weresuccessfully developed and the correlation coefficients of all models were higher than 0.9. Furthermore,the improved models with spectra preprocessed and efficient wavelength selected were obtained and theircorrelation coefficients were between 0.95-0.99.
In conclusion, the above results contribute a lot to understand the differential tolerance mechanismamong monocotyledonous and dicotyledonous species (both the crops and weeds) to ZJ0273 stress whichare the basis of safe use of this new herbicide in field. The development of NIRS models in this studywould be helpful for further field analysis of using NIRS to monitor the growing states and biologicalproperties of oilseed rape under herbicide stress.
1、离体及活体条件下，分别研究了丙酯草醚对单双子叶作物及杂草乙酰乳酸合成酶(ALS)活力的影响。在离体条件下，丙酯草醚对ALS活力几乎无抑制效果；而在活体条件下，叶面喷施丙酯草醚对双子叶作物油菜ALS活力抑制作用最小，对单子叶作物大麦抑制作用最显著，对阔叶杂草牛繁缕和禾本科杂草看麦娘的抑制效果介于前两种作物之间。植株不同部位的ALS活力存在较大差异，越幼嫩的部位酶活力越高，因此在ALS活力分析中顶端新叶是合适的研究材料。100mg／L丙酯草醚处理下牛繁缕、油菜和大麦的不同叶位ALS活力变化趋势存在差异，牛繁缕与油菜下层叶的酶活力随处理时间延长而下降的程度比上层叶剧烈，在大麦中则相反；油菜上层叶酶活力随处理时间的延长而迅速恢复，而牛繁缕和大麦则无法恢复。三种不同类型油菜耐药性表现为，芥菜型油菜耐性最强，而白菜型油菜耐性最弱，甘蓝型油菜介于两者之间。
2、探讨了丙酯草醚处理对发芽期油菜及大麦幼苗的生长发育及根细胞活性、有丝分裂、根叶细胞形态及超微结构的影响。种子发芽期是丙酯草醚处理的敏感时期；对发芽期油菜和大麦较为安全的临界浓度为1mg／L；10mg／L以上浓度处理下油菜和大麦幼苗的生长尤其是根系的伸长和发育及根尖细胞活性等方面受到了显著抑制，且抑制效果随处理浓度的升高和处理时间的延长而增大。100mg／L处理下油菜和大麦根尖细胞有丝分裂指数均下降，有丝分裂产生的细胞数量减少造成分生区和伸长区明显缩短；另一方面处理还抑制成熟区细胞的发育，导致如油菜侧根发育明显滞后等现象。100mg／L丙酯草醚处理对油菜和大麦根细胞膜造成较大的破坏并导致胞内电解质外渗；处理还对根细胞中的线粒体外膜造成了一定程度的破坏，并存在胞内微粒体数量增加的现象(推测该除草剂的代谢活化作用可能在微粒体中进行，而线粒体中可能存在其活化后的作用靶标)；丙酯草醚处理显著影响细胞亚结构的发育，如抑制大麦根细胞壁、液泡、质体及其叶细胞中的叶绿体发育，但加速油菜子叶细胞中叶绿体的衰老(推测丙酯草醚具有两重性——加速成熟组织的衰老和延缓幼嫩组织的发育，故可尝试将其开发为一种新型的生长调节剂)。在生化检测及透射电镜(TEM)观察都未发现100mg／L丙酯草醚处理对油菜及大麦根尖细胞的核膜造成明显影响，压片法观察分裂细胞也未发现由处理引起的染色体异常和细胞分裂畸形现象。
3、研究了营养生长期油菜及大麦在丙酯草醚胁迫下植株生物量积累、抗氧化系统和光合系统的差异表现及生理机制。发芽期油菜对丙酯草醚的耐性并不高于大麦，但5叶期油菜植株对丙酯草醚的耐性显著高于大麦，表现在：100 mg／L处理浓度对油菜各生理参数均无显著影响，高浓度处理对油菜植株造成显著影响但抑制作用随时间的延长而减弱：而5叶期大麦对100 mg／L处理浓度已敏感，且抑制效果随处理浓度的升高和时间的延长而加剧。在丙酯草醚胁迫下，大麦过氧化物酶(POD)和超氧化物歧化酶(SOD)活性先显著升高后迅速下降，而油菜却未表现出明显的变化趋势，可见丙酯草醚对大麦的抗氧化系统造成的影响大于油菜；在光合系统方面，丙酯草醚对油菜净光合速率(Pn)几乎无影响但却导致大麦叶片SPAD值和Pn的显著下降，这可能与大麦光合系统Ⅱ(PSⅡ)的非光化学耗散能力(NPQ)受到较大抑制有关；丙酯草醚对大麦光合系统造成较大破坏最终导致其更显著地影响大麦的生物量积累。
4、分析了苗期和抽薹期分别喷施丙酯草醚对油菜产量和种子品质等的影响。明确了在苗期喷施100mg／L丙酯草醚可促进油菜增产且对种子品质无显著影响，但1000mg／L高浓度处理对产量和品质(尤其是蛋白质积累方面)有明显抑制；而油菜抽薹期则是对丙酯草醚处理的敏感时期，即使是100mg／L田间推荐浓度喷施处理也会造成减产并影响种子品质，且处理对产量的影响比对品质的影响更显著，1000mg／L高浓度处理几乎导致颗粒无收。以上不同时期作物间的敏感性差异一方面与作物本身对丙酯草醚的活化和相关代谢途径、能力等存在差异，另一方面也可能与处理方式(根部添加和叶面喷施处理)以及丙酯草醚以根茎吸收为主的特性等原因有关。因此在生产上，作物和杂草的苗龄控制及田间施用方式等都将对丙酯草醚的除草效果及对作物的安全性等方面造成影响，应引起足够重视。
5、将近红外光谱技术用于丙酯草醚胁迫下油菜不同叶位生理信息的监测。成功地对油菜叶片中ALS活力，可溶性蛋白、非可溶性蛋白、总蛋白含量和总氨基酸含量等生理参数进行了光谱建模，模型的预测相关系数皆高于0．9：通过不同光谱预处理、有效波长选择和建模方法筛选等技术手段构建的改良模型预测相关系数高达0．95～0．99。
本研究取得的创新性结果对丙酯草醚对单、双子叶植物抑制机制差异有了更全面深入的认识，这将为该新型油菜田除草剂对油菜及后茬作物大麦等安全性评价方面提供重要参考。本研究中对丙酯草醚处理下油菜叶片的光谱数据与不同生理参数进行建模的尝试，为近红外光谱技术应用于除草剂等各种胁迫下植物药害等的大田活体无损快速监测奠定了较好基础，具有显著的创新性和实践应用意义。
Propyl 4-(2-(4,6-dimethoxypyrimidin-2-yloxy)benzylamino)benzoate (ZJ0273) is a novel herbicideused in the rapeseed field with the advantage of broad weeding spectrum. It is meaningful to study themode of action and tolerance mechanism of this new herbicide among crops and weeds for its extendedapplication. In this study, rapeseed (Brassica napus L. cv. ZS758) and barley (Hordeum vulgare L. cv.Huangqingluo) were investigated to clarify the tolerance mechanism within monocotyledon anddicotyledon species to ZJ0273 stress with various physiological measurements and microscopic analyses.In addition, several near-infrared spectroscopy (NIRS) models were developed to investigate for rapid andnon-destructive determination of physiological characters in rapeseed leaves under herbicide ZJ0273stress. The main results are as follows:
1. The differential tolerance or susceptibility of four plant species viz. B. napus L. (rapeseed),Hordeum vulgare L. (barley), Malachium aquaticum (L.) Fries (water chickweed) and Alopecurusaequalis Sobol. (equal alopecurus), to various concentrations of ZJ0273, was studied based on the rate ofinhibition of acetolactate synthase (ALS) activity. B. napus was observed as the most tolerant among fourspecies in which ALS activity in vivo was hardly affected by ZJ0273 at 100 mg/L (recommended dose inthe rapeseed field). In contrast, Hordeum vulgare was found to be more susceptible while the two weedspecies exhibited moderate susceptibility as compared to Hordeum vulgare. However, ALS activity in vitrowas hardly inhibited by herbicide ZJ0273. In addition, there was a wide variation noted for in vivo ALSactivity among various parts of plants. The uppermost younger leaf seemed to be the most appropriatesample for evaluation, as it showed the highest level of ALS activity in the plant. Spatial and temporalvariations in ALS activity as affected by 100 mg/L ZJ0273 in water chickweed, rapeseed and barley alsosuggested that the synchronized sampling based on the tissue position is necessary to examine precisely,the difference in tolerance or susceptibility to ALS-inhibiting herbicide. The variant tendency of ALSactivity from different parts of plants within monocotyledons and dicotyledons to ZJ0273 treatmentsshowed differential response which revealed some possible mechanisms related to the altered absorptionand translocation of the herbicide. Furthermore, the tolerance among three species of Brassica suggestedthat B. rapa was the most susceptible and B. juncea was the most tolerant to the new herbicide ZJ0273.
2. Response of seedling growth, root cell viability, mitosis, root and leaf cell structure andultrastructure of rapeseed and barley to herbicide ZJ0273 at germination stage was studied to define the action mechanism of this novel herbicide. The results indicated that both of rapeseed and barley seedlingsare very sensitive to ZJ0273 at germination stage and the application of 1 mg/L (optimum concentration) issafer for seedling growth. 10 mg/L ZJ0273 treatment can significantly inhibit seedling growth (especiallythe root elongation and development) and root viability. Besides, the inhibitive effects become intensifiedwith the increase of dosage and duration. Mitotic index of root-tip cells was declined and cell division wasstopped at metaphase after being treated by 100 mg/L ZJ0273, which shortened the meristematic zone andelongation zone in plant root tips. In addition, the cell development in maturation zone was also inhibited,which deferred the development of lateral root. ZJ0273 significantly inhibited the development of cell wall,vacuole, plastid and chloroplast in barley cells while induced the senescence (aging) of chloroplast inrapeseed cotyledon cells, which suggested that the effect of ZJ0273 on cell development is bifacial and itcould be exploited as a plant growth regulator. Furthermore, 100 mg/L ZJ0273 treatment obviouslyinhibited root cell viability and resulted in disintegration of cell and mitochondria membrane, as well asmore microbodies were observed in treated cells, which suggested that ZJ0273 might be activated withinthe microbodies and acting in mitochondria in plant cells. However, nucleus in root and leaf cells ofrapeseed and barley treated by ZJ0273 showed no significant change in comparison to that of control, asshown by the histochemical and transmission electron microscopy (TEM) demonstrations. Furthermore,chromosome number and shape in dividing cells treated by 100 mg/L ZJ0273 were also normal, whichindicated that this new herbicide hardly induced the tolerant weed mutants and it is helpful to itsapplication and commercialization in fields.
3. The tolerance mechanism of rapeseed and barley to ZJ0273 stress was studied by measurements ofbiomass, antioxidants and photosynthetic characteristics. As above results shows, rapeseed is no moretolerant than barley at germination stage. In contrast, the tolerance of rapeseed seedlings at 5-leaf stage issignificantly higher than barley seedlings to ZJ0273, which showed that rapeseed seedlings were hardlyaffected by 100 mg/L treatment where as the inhibitive effect of ZJ0273 treatments at higherconcentrations on physiological characteristics recovered with the passage of time. However, barley wasmore susceptible which was reflected in lower biomass production even under at the lowest rate of 100mg/L of ZJ0273 and the enhanced inhibitive effect along with the increase of treatment concentrations anddurations. In the beginning, rapid increase and then sudden decline of leaf peroxidase (POD) andsuperoxide dismutase (SOD) activities under herbicide stress, indicated that ZJ0273 affected barleyantioxidant system more badly than rapeseed, as compared to the slight changes in rapeseed. In addition,ZJ0273 treatments hardly affected net photosynthesis (Pn) of rapeseed leaves but significantly inhibitedSPAD and Pn values of barley leaves, which indicated that the photosystemⅡof barley was paralyzed byinhibition effect of herbicide on non-photochemical quenching (NPQ) capability and finally, the biomass was reduced more significantly.
4. Effect of herbicide ZJ0273 on plant agronomic characters, seed yield and quality of oilseed rapewas also compared in this study by treating at seedling stage and bolting stage, respectively. The resultsconfirmed that 100 mg/L ZJ0273 (recommended dose in the rapeseed field) promoted seed yield withslight effect on seed quality but significantly reduced rapeseed yield and affected seed quality whenapplied at seedling stage and bolting stage, respectively. The treatment at highest concentration of 1000mg/L applied at seedling stage mainly affected seed quality but badly reduced seed yield when was appliedat bolting stage, which indicated that rapeseed yield and seed quality are more sensitive to ZJ0273 atbolting stage than seedling stage. These results suggested that crop tolerance revealed some possiblemechanisms related to the altered absorption, translocation of the herbicide. In addition, treatmentapproach such as root application or leaf-spray was also possible reason that affected the herbicideinhibitive action according to its root-absorption trait. Thus, both of optimal leaf stage and treatmentapproach should be considered in field application of ZJ0273.
5. Near-infrared spectroscopy (NIRS) was investigated for rapid and non-destructive determination ofvarious physiological characters in rapeseed leaves from different leaf-positions under herbicide stress.NIRS models of ALS activity, soluble, non-soluble, total protein and total amino acid contents weresuccessfully developed and the correlation coefficients of all models were higher than 0.9. Furthermore,the improved models with spectra preprocessed and efficient wavelength selected were obtained and theircorrelation coefficients were between 0.95-0.99.
In conclusion, the above results contribute a lot to understand the differential tolerance mechanismamong monocotyledonous and dicotyledonous species (both the crops and weeds) to ZJ0273 stress whichare the basis of safe use of this new herbicide in field. The development of NIRS models in this studywould be helpful for further field analysis of using NIRS to monitor the growing states and biologicalproperties of oilseed rape under herbicide stress.
引文
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