高兴祥^,, 李健^,, 张悦丽, 李美^,, 房锋山东省农业科学院植物保护研究所山东省植物病毒学重点实验室,济南 250100

Resistance Level, Mechanism of Alopecurus myosuroides and Control Efficacy in Wheat Field in Shandong Province

GAO XingXiang^,, LI Jian^,, ZHANG YueLi, LI Mei^,, FANG FengShandong Key Laboratory of Plant Virology, Institute of Plant Protection, Shandong Academy of Agricultural Sciences, Ji’nan 250100

通讯作者: 李美,E-mail：limei9909@163.com

高兴祥和李健为同等贡献作者。
责任编辑: 岳梅
收稿日期:2020-01-16接受日期:2020-03-27网络出版日期:2020-09-01

基金资助:

国家重点研发计划.2017YFD0201700 山东省农业科学院农业科技创新工程.CXGC2018E04

Received:2020-01-16Accepted:2020-03-27Online:2020-09-01
作者简介 About authors
高兴祥,E-mail：xingxiang02@163.com。

李健,E-mail：lijian910@163.com。






摘要
【背景】大穗看麦娘（Alopecurus myosuroides）是我国大陆近几年新发展蔓延的一种恶性禾本科杂草,目前已在山东、河南、河北、安徽等省有分布,且分布面积不断扩大。【目的】明确山东省冬小麦田大穗看麦娘对常用除草剂的抗性水平及部分种群产生抗性的机理,并评价不同除草剂对其田间防除效果,为制定小麦田大穗看麦娘防控技术规程提供理论依据。【方法】室内采用整株生物测定法测定9个大穗看麦娘种群对乙酰乳酸合成酶（ALS）抑制剂除草剂啶磺草胺、甲基二磺隆以及乙酰辅酶A羧化酶（ACC）抑制剂除草剂唑啉草酯、炔草酯、精噁唑禾草灵共5种除草剂的抗性水平,并对产生抗性的种群进行靶标基因检测,同时分别在冬前和冬后开展田间试验,评价不同除草剂对大穗看麦娘的田间防除效果。【结果】室内测定结果表明,种群8（JN2）对啶磺草胺和甲基二磺隆产生明显抗性,相对抗性指数分别达到47.32、15.97,靶标基因检测显示,该种群内植株ALS基因编码的第197位点氨基酸发生由脯氨酸（CCC）到苏氨酸（ACC）的突变;所有种群对唑啉草酯、炔草酯和精噁唑禾草灵均表现敏感。田间试验结果表明,冬前使用除草剂对大穗看麦娘的防除效果优于冬后使用除草剂的效果,冬前唑啉草酯对大穗看麦娘鲜重防效为98.6%,而冬后处理为89.1%;冬前使用啶磺草胺、甲基二磺隆对大穗看麦娘的株防效和鲜重防效在72.2%—89.3%,冬后使用则为68.6%—83.2%;唑啉草酯、炔草酯和精噁唑禾草灵对大穗看麦娘均表现出很好的防除效果,冬前使用的株防效和鲜重防效均在96.2%以上,冬后使用在82.6%—92.2%。【结论】供试的9个大穗看麦娘种群中,发现1个种群对甲基二磺隆、啶磺草胺产生较高抗性,但未发现对唑啉草酯等产生抗性的种群,室内试验和田间试验结果具有一致性。
关键词： 大穗看麦娘;ALS抑制剂;ACC抑制剂;抗性水平;抗性机理

Abstract
【Background】As a new kind of malignant gramineous in the mainland of China in recent years, Alopecurus myosuroides has been widely distributed in Shandong, Henan, Hebei and Anhui provinces, and its distribution area is constantly expanding.【Objective】The objective of this study is to determine the resistance level, mechanism of A. myosuroides to herbicides and the control efficacy in wheat field, and to provide a theoretical basis for A. myosuroides control in winter wheat field.【Method】Whole-plant dose response experiments were conducted to determine the resistance level of nine populations to ALS inhibitor such as pyroxsulam, mesosulfuron-methyl and ACC inhibitor such as pinoxaden, clodinafop-propargyl and fenoxaprop-P-ethyl in the greenhouse, and the target gene of resistant population was detected. At the same time, herbicide control experiments were carried out before and after winter in the wheat field.【Result】The results of greenhouse experiment showed that only one of the nine populations, population 8 (JN2), had obvious resistance to the herbicides of ALS inhibitor (pyroxsulam and mesosulfuron-methyl), and the resistance index (RI) reached 47.32 and 15.97, respectively. ALS gene sequence analysis showed that Pro (CCC) to Thr (ACC) mutation at position 197 of the ALS gene occurred in the resistant population. Meanwhile, all the populations had no resistance to the ACC inhibitor pinoxaden, clodinafop-propargyl and fenoxaprop-P-ethyl. The results of field trials showed that the effect of application before winter was better than that after winter, the fresh weight control efficacy of pinoxaden before winter was 98.6% and that after winter was 89.1%. The plant control efficacy and fresh weight control efficacy of pyroxsulam and mesosulfuron-methyl before winter were 72.2%-89.3% and those after winter were 68.6%-83.2%, but pinoxaden, clodinafop- propargyl and fenoxaprop-P-ethyl all showed good control efficacy on A. myosuroides, the plant control efficacy and fresh weight control efficacy were above 96.2% before winter and 82.6%-92.2% after winter.【Conclusion】Among the nine tested populations, one population was found to have high resistance to mesosulfuron-methyl and pyroxsulam, but no resistant population was found to ACC inhibitor herbicides, such as pinoxaden, clodinafop-propargyl and fenoxaprop-P-ethyl. The results of laboratory test and field test were consistent.
Keywords：Alopecurus myosuroides;ALS inhibitor;ACC inhibitor;resistance level;resistance mechanism


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本文引用格式
高兴祥, 李健, 张悦丽, 李美, 房锋. 山东省小麦田大穗看麦娘抗性水平、靶标抗性机理及田间防除效果测定[J]. 中国农业科学, 2020, 53(17): 3518-3526 doi:10.3864/j.issn.0578-1752.2020.17.009
GAO XingXiang, LI Jian, ZHANG YueLi, LI Mei, FANG Feng. Resistance Level, Mechanism of Alopecurus myosuroides and Control Efficacy in Wheat Field in Shandong Province[J]. Scientia Acricultura Sinica, 2020, 53(17): 3518-3526 doi:10.3864/j.issn.0578-1752.2020.17.009

0 引言

【研究意义】大穗看麦娘（Alopecurus myosuroides）为越年生杂草,在欧洲尤其是西北欧分布广泛、危害严重、抗性问题突出,自笔者^[1]2014年首次报道大穗看麦娘在我国大陆的鲁西北平原和鲁中山区发生危害以来,目前大穗看麦娘在山东、河南、河北、安徽省等黄淮海区域冬小麦田均有分布,且在部分地区甲基二磺隆（mesosulfuron-methyl）对大穗看麦娘田间防除效果差,危害程度逐渐加重。研究大穗看麦娘对乙酰乳酸合成酶（ALS）和乙酰辅酶A羧化酶（ACC）两类除草剂的抗性水平、抗性机理,以及不同除草剂对其田间防除效果,对制定该恶性杂草区域化治理规程具有重要意义。【前人研究进展】大穗看麦娘在欧洲的分布面积最大,且在10多个国家对包括ALS抑制剂类除草剂和ACC抑制剂类除草剂在内的7种不同作用类型的除草剂产生了严重抗性^[2,3,4,5]。国外科学家对该杂草抗性机理尤其是靶标抗性方面进行了大量研究,但在国内因为是近几年发生的杂草,对其抗性水平和抗性机理报道较少。该杂草2014年首次在我国大陆发现并报道,目前已成为黄淮地区部分小麦田一种恶性杂草,且发生区域有逐渐扩散的趋势。大穗看麦娘繁殖能力强,在田间与小麦竞争水分和养分,对小麦生产造成严重威胁,房锋等^[6,7]报道大穗看麦娘出苗高峰期在小麦播后20—40 d,成熟期植株高出小麦2.5 cm,当大穗看麦娘发生密度为420株/m²时,小麦穗密度减少55.2%,穗粒数减少26.4%。国外对大穗看麦娘的报道主要集中在抗药性及治理方面,大穗看麦娘是欧洲最重要的除草剂抗性杂草之一,对精噁唑禾草灵等7种除草剂均有产生不同程度抗性的报道^[2,3],其中ACC靶标抗性广泛存在^[2,4],抗性发生主要是由靶标基因1781位点突变导致^[4,5]。对ALS类除草剂甲基二磺隆的抗性也较为普遍,主要是由ALS基因第197位点发生突变导致^[2];抗性治理主要是通过深翻、轮作以及不同作用机制除草剂混用等措施^[2]。笔者^[8]于2014年进行了甲基二磺隆、啶磺草胺（pyroxsulam）、氟唑磺隆（flucarbazone-Na）、唑啉草酯（pinoxaden）、炔草酯（clodinafop-propargyl）、精噁唑禾草灵（fenoxaprop-P-ethyl）、异丙隆（isoproturon）和肟草酮（tralkoxydim）8种药剂对大穗看麦娘的田间防除试验,结果表明啶磺草胺和甲基二磺隆无论冬前还是冬后返青初期施药均有很好的效果,唑啉草酯、炔草酯、精噁唑禾草灵冬前施药效果好。【本研究切入点】大穗看麦娘在我国发生面积相对较小,但蔓延趋势明显,且近年来市场反馈部分区域甲基二磺隆等ALS抑制剂类除草剂对大穗看麦娘防除效果下降。然而,目前国内关于该杂草对ALS类除草剂抗药性的研究报道很少,其田间治理技术也鲜有报道。【拟解决的关键问题】以山东省冬小麦田采集的9个大穗看麦娘种群为研究对象,室内测定其对甲基二磺隆、啶磺草胺和唑啉草酯、炔草酯、精噁唑禾草灵的抗性水平,并对抗性种群进行靶标基因检测;在田间通过冬前和冬后两次用药测定两类药剂对大穗看麦娘的防除效果,明确目前有效防治大穗看麦娘的药剂种类,为制定冬小麦田大穗看麦娘区域化治理对策和合理选择除草剂提供理论依据。

1 材料与方法

1.1 材料

大穗看麦娘种子：2017年5月采集于山东省有大穗看麦娘发生的6个地市（济南、德州、滨州、聊城、菏泽、临沂）冬小麦田,共采集大穗看麦娘9个种群。种群以采集地首字母命名,种群1—9分别标记为DZ-1、DZ-2（德州）、BZ-1（滨州）、LC-1（聊城）、HZ-1（菏泽）、LY-1（临沂）、JN-1、JN-2和JN-3（济南）。采集地小麦田之前主要发生的禾本科杂草为节节麦（Aegilops tauschii）和雀麦（Bromus japonicus）,施用过除草剂啶磺草胺或甲基二磺隆,未施用过唑啉草酯、炔草酯以及精噁唑禾草灵。

除草剂：4%啶磺草胺可分散油悬浮剂,美国陶氏益农公司;30 g·L^-1甲基二磺隆可分散油悬浮剂,拜耳股份公司;70%氟唑磺隆水分散粒剂,爱利思达生物化学品北美有限公司;5%唑啉草酯乳油,瑞士先正达作物保护有限公司;15%炔草酯可湿性粉剂,瑞士先正达作物保护有限公司;69 g·L^-1精噁唑禾草灵水乳剂,拜耳股份公司。

1.2 方法

1.2.1 室内整株法抗性水平测定 试验于2017年11月至2018年2月进行,将9个大穗看麦娘种群的种子分别定量播于直径为9 cm的塑料盒中,覆土1—2 mm,底部渗水至土表后,转移至玻璃温室中培养。温室自然光照,温度15—25℃。定期以盆钵底部渗灌方式补水,保持土壤湿度。

抗性测定采用盆栽整株剂量-反应测定法^[9]。啶磺草胺、甲基二磺隆剂量为0.37、1.11、3.33、10.00、30.00、90.00 g·hm^-2;唑啉草酯、炔草酯、精噁唑禾草灵剂量为0.77、2.22、6.67、20.00、60.00、180.00 g·hm^-2（以上用量均为有效成分用量）;于大穗看麦娘2叶1心期,采用ASS-4型自动控制喷洒系统茎叶均匀喷雾,用水量为450 L·hm^-2,喷雾压力0.35 MPa,扇形喷头流量800 mL·min^-1。施药后30 d称量各处理大穗看麦娘地上部分鲜重,计算鲜重抑制率。鲜重抑制率（%）=100×（空白对照鲜草重-处理鲜草重）/空白对照鲜草重。

1.2.2 抗性种群靶标基因检测 根据大穗看麦娘ALS基因序列（GenBank登录号：AJ437300.2）设计引物（AM-F 5′ GCCTTACCCAAACCTACTCT 3′和AM-R 5′ AGTTGATATAACATACACCAGCA 3′）,用于克隆靶标基因,克隆片段包含目前已报道所有抗性相关位点。待大穗看麦娘3叶期,采集叶片,提取基因组DNA,回收PCR产物并送上海生工生物工程有限公司序列测定。将测序后获得序列与敏感型进行比对,分析相关功能位点是否存在氨基酸水平突变。

1.2.3 防除大穗看麦娘田间药效测定 试验地点设在山东省济南市济阳区大邝村,小麦品种为济麦22,于2017年9月30日机械播种,浅旋。设置6种药剂处理,4次重复：4%啶磺草胺可分散油悬浮剂12 g·hm^-2、30 g·L^-1甲基二磺隆可分散油悬浮剂13.5 g·hm^-2、70%氟唑磺隆可分散粒剂36.75 g·hm^-2、5%唑啉草酯乳油52.5 g·hm^-2、15%炔草酯可湿性粉剂67.5 g·hm^-2、69 g·L^-1精噁唑禾草灵水乳剂51.75 g·hm^-2。另设空白对照处理,共计7个处理。试验1施药时间为2017年11月6日（小麦分蘖期）,试验2施药时间为2018年3月5日（小麦返青初期）。

采用绝对值（数测）调查法,每处理小区随机取4点,每点调查0.25 m²,分别记录杂草数量,冬前试验在药后30 d和冬后小麦拔节期各调查一次,冬后试验在施药后15、30 d各调查一次,最后一次在调查杂草株数的同时进行鲜重调查,计算鲜重防效。

1.3 数据分析

采用DPS 7.05软件对药剂剂量的对数值与防治效果的概率值进行回归分析,得到剂量-反应曲线、相关系数、抑制杂草生长50%的除草剂剂量（GR₅₀）及95%置信区间。以所有种群中GR₅₀值最小的种群为敏感种群,其他大穗看麦娘种群的相对抗性指数（resistance index,RI）为其GR₅₀值与敏感种群GR₅₀比值,即RI=GR₅₀（R）/GR₅₀（S）。抗性判断参考高兴祥等^[9]：1.00≤RI<5.00为敏感种群,5.00≤RI<10.00为低抗性种群,10.00≤RI<50.00为中抗性种群,RI≥50.00为高抗性种群。

2 结果

2.1 不同大穗看麦娘种群对ALS除草剂的抗性水平

9个大穗看麦娘种群中有8个种群对甲基二磺隆和啶磺草胺均敏感,药后7 d开始大穗看麦娘生长受到明显抑制,叶色黄化,逐渐死亡,但JN-2种群敏感性明显低于其他种群,对啶磺草胺、甲基二磺隆抗性明显,GR₅₀分别为16.09和11.34 g·hm^-2,按GR₅₀计算出的RI分别为47.32、15.97,防除效果均不理想,各剂量处理后JN-2种群均无死亡,仅高剂量处理有一定的生长抑制作用。8个敏感种群对甲基二磺隆、啶磺草胺GR₅₀分别在0.71—1.44、0.34—0.72 g·hm^-2,按GR₅₀计算出的RI基本一致,在1.00—2.12（表1）。

Table 1
表1
表1大穗看麦娘种群对甲基二磺隆、啶磺草胺抗性水平测定
Table 1Resistance level of A. myosuroides populations to mesosulfuron-methyl and pyroxsulam

药剂 Herbicide	种群编号 Number of population	回归方程 Regression equation (y=)	相关系数Correlation coefficient	GR50 (g·hm-2) (95% CL)	GR90 (g·hm-2) (95% CL)	相对抗性指数 RI
甲基二磺隆Mesosulfuron-methyl	1	4.8011+1.2453x	0.9756	1.44 (1.02-1.92)	15.44 (11.96-20.90)	2.03
	2	5.0663+1.1225x	0.9932	0.87 (0.54-1.26)	12.10 (9.23-16.55)	1.23
	3	4.8753+1.5563x	0.9535	1.20 (0.47-2.08)	8.01 (5.19-13.43)	1.69
	4	5.0964+1.1904x	0.9342	0.83 (0.27-1.58)	9.90 (6.10-18.34)	1.17
	5	5.2020+1.8078x	0.9292	0.77 (0.48-1.10)	3.95 (3.18-4.80)	1.08
	6	5.1410+1.0520x	0.9019	0.73 (0.43-1.10)	12.14 (9.12-16.95)	1.03
	7	5.2486+1.6669x	0.9241	0.71 (0.43-1.03)	4.17 (3.33-5.11)	1.00
	8	4.2554+0.7059x	0.9752	11.34 (7.96-17.07)	741.59 (315.44-2558.27)	15.97
	9	4.9515+1.9814x	0.9770	1.06 (0.73-1.41)	4.69 (3.90-5.58)	1.49
啶磺草胺 Pyroxsulam	1	5.2960+1.1712x	0.9327	0.56 (0.31-0.87)	6.94 (5.34-9.15)	1.65
	2	5.3536+1.1046x	0.9361	0.48 (0.25-0.77)	6.92 (5.24-9.27)	1.41
	3	5.1644+1.1325x	0.9221	0.72 (0.42-1.06)	9.69 (7.44-13.06)	2.12
	4	5.3877+1.4096x	0.9643	0.53 (0.29-0.82)	4.31 (3.33-5.45)	1.56
	5	5.3531+1.2250x	0.9413	0.51 (0.27-0.81)	5.73 (4.40-7.44)	1.50
	6	5.3454+0.7355x	0.8265	0.34 (0.14-0.63)	18.74 (12.40-32.00)	1.00
	7	5.2316+0.8877x	0.8704	0.55 (0.28-0.89)	15.23 (10.85-23.08)	1.62
	8	4.4847+0.4271x	0.9471	16.09 (9.07-35.02)	1613.87 (241.47-5545.1)	47.32
	9	6.0290+1.3518x	0.9271	0.34 (0.24-0.69)	11.54 (6.83-20.28)	1.00

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2.2 大穗看麦娘ALS除草剂抗性种群靶标基因检测

大穗看麦娘抗性种群ALS基因测序分析见图1。

图1

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图1抗性种群的突变位点分析

Fig. 1Mutation analysis of resistant population



所有检测植株均发生了ALS基因功能位点的突变,且突变方向一致,均为第197位氨基酸发生脯氨酸（CCC）到苏氨酸（ACC）的突变。该位点的突变造成乙酰乳酸合成酶结构的改变,影响与相应除草剂的结合。

2.3 不同大穗看麦娘种群对ACC除草剂的抗性水平

9个大穗看麦娘种群对唑啉草酯、炔草酯和精噁唑禾草灵均敏感,药后7 d开始大穗看麦娘生长受到明显抑制,叶色黄化,逐渐死亡,9个种群反应无明显差异。

由表2可见,9个种群对同一种药剂的敏感性无明显差异。唑啉草酯对大穗看麦娘9个种群的GR₅₀在1.64—3.15 g·hm^-2;炔草酯对9个种群的GR₅₀在1.61—2.39 g·hm^-2;精噁唑禾草灵对9个种群的GR₅₀在2.96—3.85 g·hm^-2。根据抗性结果可知,对于这3种除草剂,9个大穗看麦娘种群均属敏感种群。

Table 2
表2
表2大穗看麦娘种群对唑啉草酯、炔草酯和精噁唑禾草灵抗性水平测定
Table 2Resistance level of A. myosuroides populations to pinoxaden, clodinafop-propargyl and fenoxaprop-P-ethyl

药剂 Herbicide	种群编号 Number of population	回归方程 Regression equation (y=)	相关系数Correlation coefficient	GR50 (g·hm-2) (95% CL)	GR90 (g·hm-2) (95% CL)	相对抗性指数 RI
唑啉草酯 Pinoxaden	1	4.6163+1.2066x	0.9429	1.64 (1.01-2.38)	19.63 (15.24-26.10)	1.00
	2	4.7433+1.1897x	0.9777	1.87 (1.23-2.56)	19.11 (11.48-26.95)	1.14
	3	4.4679+1.3224x	0.9325	2.53 (1.74-3.40)	23.52 (18.61-30.79)	1.54
	4	4.4646+1.2843x	0.9502	2.61 (1.81-3.50)	26.00 (20.39-34.45)	1.59
	5	4.5855+1.3372x	0.9399	2.04 (1.35-2.82)	18.55 (14.76-24.00)	1.24
	6	4.4631+1.3000x	0.9448	2.59 (1.80-3.47)	25.05 (19.72-33.04)	1.58
	7	4.3568+1.4520x	0.9436	2.77 (1.97-3.64)	21.16 (17.06-27.00)	1.69
	8	4.3950+1.3983x	0.9624	2.71 (1.91-3.58)	22.34 (17.88-28.80)	1.65
	9	4.2807+1.4417x	0.9605	3.15 (2.30-4.09)	24.42 (19.60-31.47)	1.92
炔草酯Clodinafop-propargyl	1	4.3066+1.8352x	0.9850	2.39 (1.67-3.14)	11.92 (9.93-14.31)	1.48
	2	4.4946+1.8619x	0.9777	1.87 (1.23-2.56)	9.11 (7.48-10.95)	1.16
	3	4.5096+1.5318x	0.9777	2.09 (1.40-2.84)	14.35 (11.68-17.81)	1.30
	4	4.6245+1.3635x	0.9672	1.89 (1.22-2.63)	16.42 (13.11-20.96)	1.17
	5	4.6924+1.4791x	0.9567	1.61 (1.00-2.30)	11.87 (9.54-14.79)	1.00
	6	4.6894+1.3348x	0.9451	1.71 (1.07-2.43)	15.59 (12.39-19.95)	1.06
	7	4.5763+1.5889x	0.9668	1.85 (1.20-2.56)	11.84 (9.64-14.55)	1.15
	8	4.6349+1.6640x	0.9714	1.66 (1.04-2.33)	9.76 (7.91-11.92)	1.03
	9	4.6081+1.5914x	0.9655	1.76 (1.13-2.46)	11.26 (9.15-13.83)	1.09
精噁唑禾草灵Fenoxaprop-P-ethyl	1	4.5516+0.8188x	0.9957	3.53 (2.30-5.00)	109.64 (81.12-143.70)	1.19
	2	4.5643+0.9184x	0.9905	2.98 (1.96-4.18)	74.12 (50.63-121.29)	1.01
	3	4.4032+1.0545x	0.8808	3.68 (2.60-4.91)	60.42 (43.52-91.13)	1.24
	4	4.5766+0.8415x	0.9975	3.19 (2.06-4.52)	106.16 (68.36-190.72)	1.08
	5	4.3503+1.1587x	0.9857	3.64 (2.62-4.78)	46.42 (34.74-66.17)	1.23
	6	4.5167+0.9471x	0.9859	3.24 (2.18-4.47)	73.04 (50.41-117.56)	1.09
	7	4.4059+1.0140x	0.8687	3.85 (2.71-5.16)	70.75 (49.89-110.14)	1.30
	8	4.4943+1.0737x	0.8882	2.96 (2.03-4.02)	56.20 (33.99-67.46)	1.00
	9	4.4054+1.0744x	0.8835	3.58 (2.53-4.77)	55.75 (40.56-82.86)	1.21

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2.4 大穗看麦娘田间防控效果

2.4.1 冬前施药 药后30 d防除效果调查发现空白对照区大穗看麦娘颜色绿,生长旺盛;各药剂处理区虽然大穗看麦娘无死亡,但啶磺草胺、甲基二磺隆处理区大穗看麦娘叶片黄化,生长明显受抑制,氟唑磺隆处理区大穗看麦娘黄化程度略差;唑啉草酯、炔草酯和精噁唑禾草灵处理区大穗看麦娘心叶黄化,生长也明显受抑制,分蘖均明显少于空白对照区,6个药剂处理株防效在45.9%—69.0%,各处理间无显著差异（表3）。

Table 3
表3
表3小麦田冬前施药对大穗看麦娘田间防除效果
Table 3Weed control efficacy to A. myosuroides after 6 herbicides treated in wheat field (before winter)

药剂 Herbicide	30 d	冬后拔节期Jointing stage (2018-04-04)
	株防效Plant control efficacy (%)	株防效Plant control efficacy (%)	鲜重防效Fresh weight control efficacy (%)
甲基二磺隆Mesosulfuron-methyl	59.1±4.2aA	89.3±2.6bA	88.6±2.6bAB
啶磺草胺Pyroxsulam	56.2±3.5aA	78.4±4.1bA	72.2±2.1bB
氟唑磺隆Flucarbazone-Na	45.9±5.1aA	38.7±2.5cB	16.5±4.6cC
唑啉草酯Pinoxaden	69.0±2.6aA	98.0±0.2aA	98.6±0.2aA
炔草酯Clodinafop-propargyl	67.9±3.2aA	99.2±0.3aA	99.3±0.1aA
精噁唑禾草灵Fenoxaprop-P-ethyl	59.4±2.6aA	96.8±1.0aA	96.2±0.3aA

同列数据后不同大、小写字母分别表示经Duncan氏新复极差法检验在P<0.01和P<0.05水平差异显著。表4同
Different uppercase and lowercase letters in the same column indicate significant difference at P<0.01 and P<0.05 level by Duncan’s new multiple range test. The same as Table 4

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冬后返青初期（2018年3月4日）调查发现,啶磺草胺、甲基二磺隆以及唑啉草酯、炔草酯、精噁唑禾草灵对大穗看麦娘均有很好的防除效果,大穗看麦娘几乎全部死亡,但氟唑磺隆防除效果差,几乎无效。

冬后拔节期（2018年4月4日）调查发现,唑啉草酯、炔草酯和精噁唑禾草灵仍保持很好的效果,株防效分别为98.0%、99.2%、96.8%,鲜重防效分别为98.6%、99.3%、96.2%;啶磺草胺和甲基二磺隆处理区,少部分大穗看麦娘从基部发芽,恢复生长,株防效78.4%—89.3%,鲜重防效72.2%—88.6%,防效明显低于冬后返青初期调查结果,也明显低于唑啉草酯、炔草酯和精噁唑禾草灵的防效;氟唑磺隆效果最差,株防效和鲜重防效仅分别为38.7%、16.5%（表3）。

2.4.2 冬后施药 药后15 d杂草效果调查发现,除氟唑磺隆外,其他药剂处理对大穗看麦娘均有较好的防除效果,大穗看麦娘心叶黄化明显,生长受到抑制,部分已死亡,株防效在42.6%—61.2%,氟唑磺隆的株防效为25.2%（表4）。

Table 4
表4
表4小麦田冬后小麦返青初期施药对大穗看麦娘田间防治试验
Table 4Weed control efficacy to A. myosuroides after 6 herbicides treated in wheat field (after winter)

药剂 Herbicide	15 d	30 d
	株防效Plant control efficacy (%)	株防效Plant control efficacy (%)	鲜重防效Fresh weight control efficacy (%)
甲基二磺隆Mesosulfuron-methyl	51.2±5.2aA	83.2±2.1bB	78.2±2.8cB
啶磺草胺Pyroxsulam	49.2±4.6aA	75.2±1.6cC	68.6±3.2dB
氟唑磺隆Flucarbazone-Na	25.2±3.5bB	19.6±3.2dD	12.5±5.1eC
唑啉草酯Pinoxaden	61.2±4.1aA	92.2±1.5aA	89.1±1.1aA
炔草酯Clodinafop-propargyl	45.2±3.5aA	90.2±1.2aA	87.1±1.2aA
精噁唑禾草灵Fenoxaprop-P-ethyl	42.6±6.2aA	87.6±2.1bB	82.6±1.3bB

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药后30 d调查发现,唑啉草酯、炔草酯和精噁唑禾草灵对大穗看麦娘防除效果好,其中唑啉草酯和炔草酯防效优于精噁唑禾草灵防效,株防效分别为92.2%、90.2%、87.6%,鲜重防效分别为89.1%、87.1%、82.6%;啶磺草胺和甲基二磺隆效果略差,株防效分别为83.2%、75.2%,鲜重防效分别为78.2%、68.6%（表4）。

3 讨论

3.1 大穗看麦娘对除草剂抗性水平

冬小麦田杂草一直是制约小麦优质高产的重要因素之一,我国黄淮海冬小麦田尤其是旱茬麦田在20世纪主要以阔叶杂草为主^[10],少有禾本科杂草,进入21世纪,随着农业耕作制度变革尤其是免耕技术的推广,适合于浅土层萌发生长的禾本科杂草种类和数量均不断上升^[1],目前,对发生于旱茬小麦田的危害最为严重的节节麦和雀麦^[11,12]研究的很多,而对在我国大陆近几年快速发展的另一种禾本科杂草大穗看麦娘鲜有报道。

杂草抗性研究方面,国内****对我国小麦田播娘蒿（Descuminia sophia）^[9,13]、荠菜（Capsella bursa- pastoris） ^[14]、猪殃殃（Galium aparine）^[15,16,17]、麦家公（Lithospermum arvense）^[18]、日本看麦娘（Alopecurus japonicus）^[19]等杂草抗药性进行了大量研究,但旱茬麦田主要禾本科杂草如雀麦、节节麦抗药性鲜有报道,对潜在恶性杂草大穗看麦娘抗药性仅有2020年一篇报道^[20]。本研究室内测定结果表明,大部分大穗看麦娘种群对ALS抑制剂啶磺草胺、甲基二磺隆和ACC抑制剂唑啉草酯、炔草酯、精噁唑禾草灵5种药剂敏感,但JN-2种群已对啶磺草胺、甲基二磺隆产生明显抗性,且相对抗性指数分别达到47.32、15.97,该结果表明,虽然大穗看麦娘在我国冬小麦田发生时间不长,但是部分区域大穗看麦娘种群已经对ALS抑制剂除草剂产生明显抗性,同时供试种群中未发现对ACC抑制剂除草剂产生抗性的种群。这一结果与葛鲁安等^[20]报道的有大穗看麦娘种群对ACC抑制剂精噁唑禾草灵产生抗性有差异,这与采集大穗看麦娘植株的区域不同有关,本试验采集的大穗看麦娘均为山东省冬小麦田,而葛鲁安等采集的为河南种群。虽然种群采集地点不同,但结合国外报道也可以看出,大穗看麦娘是一种容易对这些药剂产生抗性的禾本科杂草。

3.2 大穗看麦娘抗性种群靶标基因检测

杂草对除草剂的抗药性分为靶标抗性和非靶标抗性,其中以靶标基因突变导致抗药性占主要部分。本试验对产生抗性的大穗看麦娘种群进行的靶标基因检测结果表明,该抗性种群是ALS基因197位点发生突变引起的靶标抗性,与国外****报道^[21]的ALS基因197或574位点突变有一定的一致性。本试验采集大穗看麦娘种群均是山东省冬小麦田种群,测定结果表明未对ACC类除草剂产生抗性,但国外研究显示欧洲大穗看麦娘大部分种群除对ALS抑制剂类除草剂有抗性外,对ACC抑制剂除草剂也产生抗药性,其中对ACC抑制剂类抗性来自ACC基因1781、1999、2027、2041、2078、2088及2096等某个或某几个位点的突变^[22,23],这与欧洲大穗看麦娘发生时间长,应用相关除草剂历史时间长有关;国内葛鲁安等^[20]的研究表明,河南大穗看麦娘种群2041位点突变和2096位点突变是种群对精噁唑禾草灵产生靶标抗性的原因。

3.3 除草剂对大穗看麦娘田间防除效果及治理策略

田间试验结果表明,唑啉草酯、炔草酯和精噁唑禾草灵冬前施药对大穗看麦娘均有较好的防治效果,最终鲜重防效依然可达96.0%以上,这与前期报道结果^[8]一致;啶磺草胺和甲基二磺隆冬前施药对大穗看麦娘也有较好的防治效果,但后期效果略有下降,鲜重防效在72.2%—88.6%,这一效果明显低于前期报道^[8],可能与试验地中大穗看麦娘已经对啶磺草胺和甲基二磺隆产生一定抗药性有关。综合室内试验和田间试验结果,可以推测部分大穗看麦娘种群对啶磺草胺、甲基二磺隆已经产生一定程度的抗药性,未发现对唑啉草酯、炔草酯以及精噁唑禾草灵产生抗药性的种群。

在多个欧洲国家,大穗看麦娘对7种不同作用类型的苗后除草剂均产生了抗性^[3],因此,需密切关注大穗看麦娘对小麦田不同除草剂抗性的发生和发展,针对我国不同地区小麦栽培方式、水肥管理和除草剂使用情况,提倡多种作用机制的除草剂混合使用或者不同机理的除草剂轮换使用,进行大穗看麦娘综合治理,延缓其抗药性发展^[24,25]。另外,国内外****^{[26,27,28,29]}在小麦田土壤封闭除草剂方面也进行了大量的研究工作,小麦田土壤封闭除草剂的研究实施也是延缓抗药性,进行大穗看麦娘等杂草综合防治的重要策略之一^[30,31,32]。

4 结论

在9个大穗看麦娘种群中,发现1个种群对ALS抑制剂除草剂啶磺草胺、甲基二磺隆产生明显抗性,且属于靶标抗性,为ALS基因197位点发生突变;未发现对ACC抑制剂除草剂产生抗性的大穗看麦娘种群,但国外以及我国河南已有报道对ACC类除草剂产生抗性的大穗看麦娘种群,所以应该先明确当地小麦田大穗看麦娘的抗性现状,再有针对性地推广除草剂种类。

参考文献 View Option 原文顺序
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[1] 高兴祥, 李美, 房锋, 张悦丽, 孙作文, 齐军山. 山东省小麦田杂草组成及群落特征
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A weed survey was conducted using inverted W-pattern sampling methods to determine the species composition and structure of weed communities in wheat fields in Shandong Province. The taxa found included 69 weed species belonging to 21 families and 54 genera. Among these, 10 species were considered dominant weeds, including Descurainia sophia, Capsella bursa-pastoris, Galium aparine, Bromus japonicus, Silene conoidea, Erysimum cheiranthoides, Lithospermum arvense, Alopecurus aequalis, Aegilops squarrosa, and Calystegin hederacea; 15 species were regional dominant weeds and 44 were normal weed species. Fields in the plain regions of southwest Shandong possessed highest species richness, Shannon-wiener index and Evenness index,while the highest Simpson’s index was found in the plain regions of northwest Shandong and coastal regions of north Shandong. Hierarchical cluster analysis revealed that weeds in Shandong Province fall into 4 regional groups: hill regions of east Shandong with plain and mountain regions of mid Shandong; mountain regions of south Shandong and plain regions of southwest Shandong; plain regions of northwest Shandong ; and coastal regions of north Shandong.
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Acta Prataculturae Sinica, 2014,23(5):92-98. (in Chinese)
DOI:10.11686/cyxb20140510URL [本文引用: 2]
A weed survey was conducted using inverted W-pattern sampling methods to determine the species composition and structure of weed communities in wheat fields in Shandong Province. The taxa found included 69 weed species belonging to 21 families and 54 genera. Among these, 10 species were considered dominant weeds, including Descurainia sophia, Capsella bursa-pastoris, Galium aparine, Bromus japonicus, Silene conoidea, Erysimum cheiranthoides, Lithospermum arvense, Alopecurus aequalis, Aegilops squarrosa, and Calystegin hederacea; 15 species were regional dominant weeds and 44 were normal weed species. Fields in the plain regions of southwest Shandong possessed highest species richness, Shannon-wiener index and Evenness index,while the highest Simpson’s index was found in the plain regions of northwest Shandong and coastal regions of north Shandong. Hierarchical cluster analysis revealed that weeds in Shandong Province fall into 4 regional groups: hill regions of east Shandong with plain and mountain regions of mid Shandong; mountain regions of south Shandong and plain regions of southwest Shandong; plain regions of northwest Shandong ; and coastal regions of north Shandong.

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Frontiers in Plant Science, 2019,10:837.
DOI:10.3389/fpls.2019.00837URLPMID:31297127 [本文引用: 3]
The evolution of resistance to herbicides is a classic example of rapid contemporary adaptation in the face of a novel environmental stress. Evolutionary theory predicts that selection for resistance will be accompanied by fitness trade-offs in environments where the stress is absent. Alopecurus myosuroides, an autumn-germinating grass weed of cereal crops in North-West Europe, has evolved resistance to seven herbicide modes-of-action, making this an ideal species to examine the presence and magnitudes of such fitness costs. Here, we use two contrasting A. myosuroides phenotypes derived from a common genetic background, one with enhanced metabolism resistance to a commercial formulation of the sulfonylurea (ALS) actives mesosulfuron and iodosulfuron, and the other with susceptibility to these actives (S). Comparisons of plant establishment, growth, and reproductive potential were made under conditions of intraspecific competition, interspecific competition with wheat, and over a gradient of nitrogen deprivation. Herbicide dose response assays confirmed that the two lines had contrasting resistance phenotypes, with a 20-fold difference in resistance between them. Pleiotropic effects of resistance were observed during plant development, with R plants having a greater intraspecific competitive effect and longer tiller lengths than S plants during vegetative growth, but with S plants allocating proportionally more biomass to reproductive tissues during flowering. Direct evidence of a reproductive cost of resistance was evident in the nitrogen deprivation experiment with R plants producing 27% fewer seed heads per plant, and a corresponding 23% reduction in total seed head length. However, these direct effects of resistance on fecundity were not consistent across experiments. Our results demonstrate that a resistance phenotype based on enhanced herbicide metabolism has pleiotropic impacts on plant growth, development and resource partitioning but does not support the hypothesis that resistance is associated with a consistent reproductive fitness cost in this species. Given the continued difficulties associated with unequivocally detecting costs of herbicide resistance, we advocate future studies that adopt classical evolutionary quantitative genetics approaches to determine genetic correlations between resistance and fitness-related plant life history traits.

[4] BELZ R G, FAROOQ M B, WAGER J. Does selective hormesis impact herbicide resistance evolution in weeds? ACCase-resistant populations of Alopecurus myosuroides Huds
a, 2018,74(8):1880-1891.
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[5] KESHTKAR E, MATHIASSEN S K, MOSS S R, KUDSK P. Resistance profile of herbicide-resistant Alopecurus myosuroides (black-grass) populations in Denmark
Crop Protection, 2015,69:83-89.
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植物保护学报, 2018,45(2):340-346.
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FANG F, LI M, GAO X X, LI J, LI Y, Lü S H. Population dynamics of black-grass Alopecurus myosuroides in wheat fields and its effect on wheat yield components
Journal of Plant Protection, 2018,45(2):340-346. (in Chinese)
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[7] 房锋, 魏代平, 李美, 高兴祥, 李健, 李燕. 不同密度大穗看麦娘对小麦产量的影响
山东农业科学, 2017,49(8):119-123.
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FANG F, WEI D P, LI M, GAO X X, LI J, LI Y. Effects of different densities of Alopecurus myosuroides on wheat yield
Shandong Agricultural Sciences, 2017,49(8):119-123. (in Chinese)
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[8] 高兴祥, 孙作文, 李美, 房锋, 李健, 吴加军, 李茂昇. 大穗看麦娘化学防控田间效果评价
草业学报, 2016,25(8):172-179.
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GAO X X, SUN Z W, LI M, FANG F, LI J, WU J J, LI M S. The effect of herbicide application timing and dosage on control of Alopecurus myosuroides in winter wheat
Acta Prataculturae Sinica, 2016,25(8):172-179. (in Chinese)
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[9] 高兴祥, 李美, 高宗军, 房锋, 张悦丽, 齐军山. 山东省小麦田播娘蒿对苯磺隆的抗性测定
植物保护学报, 2014,41(3):373-378.
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GAO X X, LI M, GAO Z J, FANG F, ZHANG Y L, QI J S. Determination of flixweed (Descurainia sophia) resistance to tribenuron-methyl in Shandong Province
Journal of Plant Protection, 2014,41(3):373-378. (in Chinese)
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[10] 王金信. 山东省麦田杂草发生及其化学防除
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WANG J X. The weed occurrence and chemical control in wheat field in Shandong Province
Pesticides, 1998,37(2):11-12, 19. (in Chinese)
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[11] 房锋, 高兴祥, 魏守辉, 李燕, 李美, 张朝贤. 麦田恶性杂草节节麦在中国的发生发展
草业学报, 2015,24(2):194-201.
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FANG F, GAO X X, WEI S H, LI Y, LI M, ZHANG C X. Occurrence and effects of Aegilops tauschii in China
Acta Prataculturae Sinica, 2015,24(2):194-201. (in Chinese)
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[12] 高兴祥, 张悦丽, 李美, 房锋, 李健. 节节麦等三种禾本科杂草对不同性状土壤的适应性
植物保护学报, 2019,46(4):832-839.
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GAO X X, ZHANG Y L, LI M, FANG F, LI J. Adaptability of gramineous weeds Aegilops tauschii, Alopecurus myosuroides and Lolium multiflorum to different types of soil
Journal of Plant Protection, 2019,46(4):832-839. (in Chinese)
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[13] 高兴祥, 张悦丽, 李美, 李健, 房锋. 山东省小麦田播娘蒿对双氟磺草胺抗性水平及靶标抗性机理
中国农业科学, 2020,53(12):2399-2409.
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GAO X X, ZHANG Y L, LI M, LI J, FANG F. Resistance level and mechanism of Descurainia sophia to florasulam in wheat field of Shandong Province
Scientia Agricultura Sinica, 2020,53(12):2399-2409. (in Chinese)
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[14] 刘君良, 王金信, 刘伟堂, 金涛, 李小芳, 毕亚玲. 中国北方部分地区麦田荠菜对苯磺隆的抗性水平
农药学学报, 2011,13(4):347-353.
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LIU J L, WANG J X, LIU W T, JIN T, LI X F, BI Y L. Resistance level of Capsella bursa-pastoris to tribenuron-methyl in winter wheat fields in northern China
Chinese Journal of Pesticide Science, 2011,13(4):347-353. (in Chinese)
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[15] 马鹏生. 猪殃殃抗双氟磺草胺的生理机制研究
山东农业大学学报(自然科学版), 2014,45(5):778-781, 784.
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MA P S. Study on the physiological mechanism of the resistance of Galium aparine L. against florasulam
Journal of Shandong Agricultural University (Natural Science Edition), 2014,45(5):778-781, 784. (in Chinese)
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[16] 彭学岗, 王金信, 段敏, 杨纪辉. 中国北方部分冬麦区猪殃殃对苯磺隆的抗性水平
植物保护学报, 2008,35(5):458-462.
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PENG X G, WANG J X, DUAN M, YANG J H. The resistance to tribenuron-methyl in Galium aparine in winter wheat fields in northern China
Journal of Plant Protection, 2008,35(5):458-462. (in Chinese)
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[17] 孙健, 王金信, 张宏军, 刘君良, 卞圣楠. 抗苯磺隆猪殃殃乙酰乳酸合成酶的突变研究
中国农业科学, 2010,43(5):972-977.
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SUN J, WANG J X, ZHANG H J, LIU J L, BIAN S N. Study on mutations in ALS of resistance to tribenuron-methyl in Galium aparine L
Scientia Agricultura Sinica, 2010,43(5):972-977. (in Chinese)
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[18] 吴小虎, 王金信, 刘伟堂, 郭鹤久, 崔夕英, 陈业兵. 山东省部分市县麦田杂草麦家公Lithospermum arvense对苯磺隆的抗药性
农药学学报, 2011,13(6):597-602.
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WU X H, WANG J X, LIU W T, GUO H J, CUI X Y, CHEN Y B. Resistance of Lithospermum arvense to tribenuron-methyl in winter wheat fields in part of Shandong Province
Chinese Journal of Pesticide Science, 2011,13(6):597-602. (in Chinese)
[本文引用: 1]

[19] 李蓉荣, 袁国徽, 郭文磊, 赵祖英, 毕亚玲, 王金信. 八种除草剂对小麦田三种抗精噁唑禾草灵杂草的生物活性
植物保护学报, 2016,43(3):507-513.
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LI R R, YUAN G H, GUO W L, ZHAO Z Y, BI Y L, WANG J X. Biological activities of eight herbicides to three species of fenoxaprop-P-resistant weeds in wheat fields
Journal of Plant Protection, 2016,43(3):507-513. (in Chinese)
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[20] 葛鲁安, 朱宝林, 赵宁, 赵孔平, 王恒智, 刘伟堂, 王金信. 小麦田大穗看麦娘对精噁唑禾草灵的抗性
农药学学报, 2020,22(1):82-87.
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GE L A, ZHU B L, ZHAO N, ZHAO K P, WANG H Z, LIU W T, WANG J X. Resistance of Alopecurus myosuroides in wheat against fenoxaprop-P-ethyl
Chinese Journal of Pesticide Science, 2020,22(1):82-87. (in Chinese)
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[21] GARDIN J A C, GOUZY J, CARRèRE S, DéLYE C. ALOMYbase, a resource to investigate non-target-site-based resistance to herbicides inhibiting acetolactate-synthase (ALS) in the major grass weed Alopecurus myosuroides (black-grass)
BMC Genomics, 2015,16(1):590.
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[22] BAILLY G C, DALE R P, ARCHER S A, WRIGHT D J, KAUNDUN S S. Role of residual herbicides for the management of multiple herbicide resistance to ACCase and ALS inhibitors in a black-grass population
Crop Protection, 2012,34:96-103.
[本文引用: 1]

[23] KESHTKAR E, MATHIASSEN S K, AGHAALIKHANI M, KUDSK P. Differences in growth, development and innate seed dormancy of susceptible and fenoxaprop-P non-target site resistant black-grass sub-populations
Crop Protection, 2020, 129: Article 105022.
DOI:10.1016/j.cropro.2006.09.016URLPMID:32287715 [本文引用: 1]
Carnation mottle virus (CarMV), Carnation etched ring virus (CERV), Carnation vein mottle virus (CVMV), Carnation ringspot virus (CRSV), Carnation Italian ringspot virus (CIRV) and Carnation latent virus (CLV) are the most important viruses affecting carnation crops. All except CERV are RNA viruses. Viral RNA or DNA accumulation on root, stem, leaf, sepal, petal, stamen, pistil and ovary tissues of infected carnation or Saponaria vaccaria plants was analysed by non-isotopic molecular hybridisation. High-titres of CarMV, CRSV, CIRV, and CLV accumulated in all plant tissues whereas CERV and CVMV were irregularly distributed over the plant. High-titres of all viruses accumulated in leaf, petal, stamen, pistil, and ovary tissues, so leaves or petals are a good tissue for routine diagnosis. Six chemicals were evaluated for inactivation of all carnation viruses in infected extracts. Commercial bleach at 7% (v/v) or NaOH at 0.5% (w/v) was found to inactivate all viruses after 60 s treatment in a systemic S. vaccaria bioassay.

[24] 李美, 高兴祥, 李健, 房锋, 孙作文. 黄淮海冬小麦田杂草发生现状、防除难点及防控技术
山东农业科学, 2016,48(11):119-124.
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LI M, GAO X X, LI J, FANG F, SUN Z W. Occurrence status, control difficulties and control techniques of weeds in winter wheat field in Huang-Huai-Hai region
Shandong Agricultural Science, 2016,48(11):119-124. (in Chinese)
[本文引用: 1]

[25] 高兴祥, 李尚友, 李美, 李恩福, 李健, 房锋. 土层深度对三种麦田禾本科杂草出苗及生长的影响
植物保护学报, 2019,46(5):1132-1137.
[本文引用: 1]

GAO X X, LI S Y, LI M, LI E F, LI J, FANG F. Effects of soil depths on seedling emergence and growth of three gramineous weeds in wheat fields
Journal of Plant Protection, 2019,46(5):1132-1137. (in Chinese)
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[26] DüCKER R, Z?LLNER P, PARCHARIDOU E, RIES S, LORENTZ L, BEFFA R. Enhanced metabolism causes reduced flufenacet sensitivity in black-grass ( Alopecurus myosuroides Huds.) field populations
Pest Management Science, 2019,75(11):2996-3004.
DOI:10.1002/ps.5414URLPMID:30891919 [本文引用: 1]
BACKGROUND: Black-grass (Alopecurus myosuroides Huds.) is a frequent grass weed that commonly occurs in winter wheat in temperate Europe. Evolving resistance to post-emergence herbicides, e.g. acetyl CoA carboxylase (ACCase) and acetolactate synthase (ALS) inhibitors requires more complex weed management strategies and ensuring good efficacy of pre-emergence treatments becomes increasingly important. Flufenacet, in particular, has become a key herbicide for the control of multiple-resistant A. myosuroides. However, in some of those populations, reduced flufenacet efficacy was already observed. RESULTS: In a screening of black-grass populations from several European countries, most populations were controlled with the registered field rate of flufenacet. However, differences in the level of flufenacet sensitivity were observed and correlated with glutathione S-transferase-mediated enhanced flufenacet metabolism. The efficacy of the pre-emergence herbicides pendimethalin, prosulfocarb, S-metolachlor and pethoxamid, was also significantly decreased in populations with reduced flufenacet sensitivity. The use of flufenacet in mixtures with diflufenican, particularly in combination with flurtamone or metribuzin, however, significantly improved efficacy in less susceptible black-grass populations. CONCLUSIONS: In several populations of different European origins, reduced efficacy of flufenacet was observed due to enhanced metabolism. Although differences between populations were relatively small, best weed management practices (e.g. application of full dose rates and herbicide mixtures and wide crop rotations) should be applied to reduce selection pressure and prevent flufenacet resistance from further evolving. This is particularly important as flufenacet is one of the few still-effective herbicides suitable for the control of multiple-resistant A. myosuroides genotypes in Europe, whereas alternative pre-emergence herbicides were less effective against multiple-resistant A. myosuroides populations. (c) 2019 Society of Chemical Industry.

[27] 高兴祥, 李美, 孙作文, 房锋, 李健. 氟噻草胺对冬小麦田杂草生物活性及小麦安全性的室内测定
植物保护学报, 2017,44(5):879-880.
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GAO X X, LI M, SUN Z W, FANG F, LI J. Weed control effects of flufenacet and its safety to wheat in glasshouses
Journal of Plant Protection, 2017,44(5):879-880. (in Chinese)
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[28] 徐洪乐, 樊金星, 张宏军, 吴仁海, 苏旺苍, 薛飞, 孙兰兰, 鲁传涛. 麦田新型除草剂砜吡草唑的除草活性
植物保护, 2019,45(4):288-292.
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XU H L, FAN J X, ZHANG H J, WU R H, SU W C, XUE F, SUN L L, LU C T. Herbicidal activity of a novel herbicide pyroxasulfone in wheat field
Plant Protection, 2019,45(4):288-292. (in Chinese)
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[29] 高兴祥, 李建勇, 李美, 房锋, 李健, 齐军山. 吡氟酰草胺除草活性及对小麦安全性测定
植物保护学报, 2016,43(2):329-335.
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GAO X X, LI J Y, LI M, FANG F, LI J, QI J S. Weed control effect of diflufenican and its safety to wheat in glasshouses
Journal of Plant Protection, 2016,43(2):329-335. (in Chinese)
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[30] METCALFE H, MILNE A E, HULL R, MURDCH A J, STORKEY J. The implications of spatially variable pre-emergence herbicide efficacy for weed management
Pest Management Science, 2018,74(3):755-765.
DOI:10.1002/ps.4784URLPMID:29095563 [本文引用: 1]
BACKGROUND: The efficacy of pre-emergence herbicides within fields is spatially variable as a consequence of soil heterogeneity. We quantified the effect of soil organic matter on the efficacy of two pre-emergence herbicides, flufenacet and pendimethalin, against Alopecurus myosuroides and investigated the implications of variation in organic matter for weed management using a crop-weed competition model. RESULTS: Soil organic matter played a critical role in determining the level of control achieved. The high organic matter soil had more surviving weeds with higher biomass than the low organic matter soil. In the absence of competition, surviving plants recovered to produce the same amount of seed as if no herbicide had been applied. The competition model predicted that weeds surviving pre-emergence herbicides could compensate for sublethal effects even when competing with the crop. The ED50 (median effective dose) was higher for weed seed production than seedling mortality or biomass. This difference was greatest on high organic matter soil. CONCLUSION: These results show that the application rate of herbicides should be adjusted to account for within-field variation in soil organic matter. The results from the modelling emphasised the importance of crop competition in limiting the capacity of weeds surviving pre-emergence herbicides to compensate and replenish the seedbank. (c) 2017 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

[31] COLBACH N, CHAUVEL B, DARMENCY H, DELYE C, CORRE V L. Choosing the best cropping systems to target pleiotropic effects when managing single-gene herbicide resistance in grass weeds. A blackgrass simulation study
Pest Management Science, 2016,72(10):1910-1925.
DOI:10.1002/ps.4230URLPMID:26751723 [本文引用: 1]
BACKGROUND: Managing herbicide-resistant weeds is becoming increasingly difficult. Here we adapted the weed dynamics model AlomySys to account for experimentally measured fitness costs linked to mutants of target-site resistance to acetyl-coenzyme A carboxylase (ACCase)-inhibiting herbicides in Alopecurus myosuroides. We ran simulations to test how effectively cultural practices manage resistance. RESULTS: Simulations of an oilseed rape/winter wheat/winter barley rotation showed that, when replacing one of the seven applied herbicides with an ACCase-inhibiting one, resistant mutants exceeded 1 plant m(-2) , with a probability of 40%, after an average of 18 years. This threshold was always exceeded when three or four ACCase-inhibiting herbicides were used, after an average of 8 and 6 years respectively. With reduced herbicide rates or suboptimal spraying conditions, resistance occurred 1-3 years earlier in 50% of simulations. Adding spring pea to the rotation or yearly mouldboard ploughing delayed resistance indefinitely in 90 and 60% of simulations respectively. Ploughing also modified the genetic composition of the resistant population by selecting a previously rare mutant that presented improved pre-emergent growth. The prevalence of the mutations was influenced more by their associated fitness cost or benefit than by the number of ACCase-inhibiting herbicides to which they conferred resistance. CONCLUSION: Simulations allowed us to rank weed management practices and suggest that pleiotropic effects are extremely important for understanding the frequency of herbicide resistance in the population. (c) 2016 Society of Chemical Industry.

[32] 范志业, 沈海龙, 陈琦, 刘迪, 侯艳红, 李世民. 吡氟·氟噻·呋草酮悬浮剂土壤封闭和茎叶处理对冬小麦田阔叶杂草的防除效果比较
农药, 2017,56(1):65-68.
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FAN Z Y, SHEN H L, CHEN Q, LIU D, HOU Y H, LI S M. Control effects of diflufenican·flufenacet·flurtamone 360 g/L SC on broadleaf weeds in winter wheat field with soil and foliar treatment
Agrochemicals, 2017,56(1):65-68. (in Chinese)
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