绿豆（Vigna radiata）是我国种植的主要食用豆类之一，具有生育期短、适播期长、抗旱、耐瘠薄、固氮养地等特性，在调整种植结构、旱薄山区农民增收、丰富人们膳食结构等方面具有重要作用[1]。绿豆籽粒富含多种生理活性物质，具有清热解毒、保肝明目等保健功能，是高蛋白、低脂肪、医食同源作物[2⇓-4]。随着人们健康意识的提高，对绿豆的需求显著上升。但近年来，绿豆枯萎病（Fusarium wilt）发生日趋严重。据国家食用豆产业技术体系2017-2019年调研显示，绿豆枯萎病已在我国绿豆主产区内蒙古、吉林、黑龙江、安徽、河北等地普遍发生，尤其在西北及华北地区发生严重，绿豆枯萎病系土传维管束真菌病害，病原菌为尖镰孢菌（Fusarium oxysporum）[5-6]。病菌首先侵染小根和须根，由根部分别向根冠和茎基部扩展，病部腐烂，后期侧根和主根大部分干缩，植株容易拔起，根和茎部皮层组织及维管束变褐，叶脉间褪绿变黄，叶尖和叶缘焦枯，早期侵染可导致植株严重矮化。绿豆枯萎病一般导致20%~30%的产量损失，严重可达到50%~60%，甚至绝产，绿豆枯萎病已成为制约绿豆产业发展的重要病害[7]。目前，枯萎病防治方法包括种植抗病品种、化学防治[8-9]、生物防治[10-11]、与非寄主作物进行轮作[12⇓⇓⇓-16]等。绿豆枯萎病病原菌厚垣孢子抗逆性强，在土壤中存活时间长，农业及化学防治方法均难以控制[17]。因此，筛选抗性种质、培育和应用抗性品种是解决枯萎病发生的根本方法[18⇓-20]。
1 材料与方法
1.1 试验材料
以近年来杂交选育的81份绿豆高代品系为试验材料。接种体为尖镰孢菌株F08，由中国农业科学院作物科学研究所提供。
1.2 试验设计
1.2.1 新品系产量比较试验
绿豆新品系产量比较试验于2017-2018年在河北省农林科学院粮油作物研究所藁城堤上试验站进行，前茬为冬小麦。试验按随机区组排列，3次重复，每个品种4行区，行长4.5 m，行距50 cm，株距16.7 cm，小区面积9 m2，留苗密度1.20×105株/hm2（8000株/亩）；冀绿7号为对照品种。
1.2.2 新品种区域试验
河北省绿豆新品种区域试验于2019-2020年在8个试点（石家庄、保定、唐山、廊坊、邯郸、衡水、张家口、阳原）进行，参试品种9个（含对照品种保绿942），田间试验采用随机区组排列，3次重复，4行区，行长5.0 m，行距50 cm，株距16.7 cm，小区面积10 m2；留苗密度1.20×105株/hm2（8000株/亩）。
1.2.3 生产试验
绿豆品种生产试验于2020年在5个试点（石家庄、保定、唐山、廊坊、张家口）进行，参试品种2个，以保绿942为对照品种。试验不设重复，每个品种种植面积333.3 m2（0.5亩），行距50 cm，株距16.7 cm，留苗密度1.20×105株/hm2（8000株/亩）。
1.3 试验方法
1.3.1 苗期枯萎病抗性鉴定方法
将绿豆种子播种于以蛭石为基质的纸杯中，每杯播种1份材料，15粒，重复3次，播种后放于25 ℃的温室中培养，根据实际情况适量浇水，待幼苗2片真叶完全展开后，选取长势良好、整齐、健壮的10株幼苗进行接种，抖落幼苗根部蛭石，用自来水洗净根部，剪去约1/3根系，将根系浸泡在浓度为1×106个/mL的孢子悬浮液中约5 min，随后移栽至新的以蛭石为基质的纸杯中，接种后的幼苗置于25 ℃恒温培养室，12 h光照/12 h黑暗交替条件下培养，接菌后第14天调查植株发病情况，并计算病情指数。
1.3.2 病害分级标准
参照朱琳等[21]的绿豆尖镰孢枯萎病0~4分级标准。0级：叶片无症状；1级；叶片边缘或叶尖轻微褪绿变黄，黄化面积≤25%；2级：25%<叶片黄化面积≤50%；3级：50%<叶片重度黄化或萎蔫面积≤75%；4级：叶片完全枯萎或黄化面积>75%。
1.3.3 病情指数及抗性评价标准
苗期抗性评价依据病情指数调查判定抗性水平。
病情指数（disease index，DI）=∑（各级病叶数×各级代表值）/（调查总叶数×最高级代表值）× 100。
按病情指数划分抗性评价。高抗（HR）：0<DI≤15；抗（R）：15<DI≤35；中抗（MR）：35<DI≤55；感（S）：55<DI≤75；高感（HS）：DI>75。
成熟期抗性评价依据茎斜截面变色程度，标准参考Ulloa等[23]的方法，0级（HR）：茎斜截面无变色；1级（R）：茎斜截面出现轻度变色；2级（MR）：茎斜截面1/4~1/2区域出现变色，颜色较浅；3级（S）：大部分维管组织变色，颜色中度；4级（HS）：整个茎斜截面变色，颜色较深，植株严重受损。
1.4 数据处理
采用Excel整理并分析数据。
2 结果与分析
2.1 绿豆新品系苗期枯萎病抗性鉴定
图1
图1
绿豆新品系抗病和感病材料接种尖镰孢菌14 d后表型
Fig.1
Phenotypes after 14 days of inoculation of disease-resistant and disease-susceptible materials of new strains of mung bean with F.oxysporum
表1 绿豆新品系苗期枯萎病抗性鉴定
Table 1
| 序号 Number | 品系 Line | 病情指数 DI | 抗性评价 Resistance evaluation | 序号 Number | 品系 Line | 病情指数 DI | 抗性评价 Resistance evaluation |
|---|---|---|---|---|---|---|---|
| 1 | HN0913-6-6-1-4-4-1 | 4.47 | HR | 8 | 1105-12 | 18.02 | R |
| 2 | HN0913-6-6-1-4-1-1 | 11.43 | HR | 9 | HN0802-1-2-2-1-1-4 | 26.55 | R |
| 3 | 0914-13-1-2 | 4.93 | HR | 10 | HN0913-6-10-3-2-1-3 | 29.50 | R |
| 4 | 1111反-1 | 5.91 | HR | 11 | 1015-4-1-1 | 31.59 | R |
| 5 | HN1012-5-4-2 | 9.60 | HR | 12 | 1110反-3 | 33.10 | R |
| 6 | HN1012-6-2-2 | 14.81 | HR | 13 | 1014反-2-3-2 | 33.47 | R |
| 7 | 1105-7 | 16.64 | R | 14 | 1109-8 | 34.72 | R |
2.2 抗性品系农艺性状表现
2.2.1 绿豆新品系高级产量比较试验
14份抗病品系参加了2018年绿豆新品系高级产量比较鉴定试验，田间调查了参试品系的田间枯萎病抗性、生育期、株高、单株分枝、单株结荚、单荚粒数等主要农艺性状，并对产量进行了分析（表2）。结果表明，在14个抗性品系中，品系HN0802-1-2-2- 1-1-4和0914-13-1-2较对照种冀绿7号增产显著，产量分别为1554.00和1438.20 kg/hm2，分别增产23.75%和14.53%。其中，HN0802-1-2-2-1-1-4增产最高，且综合农艺性状和产量性状均表现突出，该品系夏播生育期65 d，株高68.0 cm，主茎分枝数4.4，单株荚数23.4，单荚粒数14.5，百粒重6.85 g，为大粒种，提交参加了2019-2020年河北省绿豆新品种区域试验。
表2 抗性品系相关农艺性状及产量表现
Table 2
| 序号 Number | 品种（系） Variety (line) | 生育期 Growth period (d) | 株高 Plant height (cm) | 单株分枝数 Branch number per plant | 单株荚数 Pods per plant | 单荚粒数 Seeds per pod | 百粒重 100-seed weight (g) | 产量 Yield (kg/hm2) | 较对照增产率 Yield increase than control (%) |
|---|---|---|---|---|---|---|---|---|---|
| 1 | HN0802-1-2-2-1-1-4 | 65 | 68.0 | 4.4 | 23.4 | 14.5 | 6.85 | 1554.00 | 23.75* |
| 2 | 0914-13-1-2 | 65 | 74.0 | 3.2 | 18.9 | 10.1 | 6.63 | 1438.20 | 14.53* |
| 3 | 1015-4-1-1 | 65 | 64.0 | 3.3 | 12.3 | 7.5 | 6.68 | 1212.60 | -3.43 |
| 4 | 1111反-1 | 65 | 74.0 | 3.8 | 19.6 | 10.4 | 6.40 | 1209.00 | -3.73 |
| 5 | 1105-7 | 65 | 70.0 | 3.9 | 16.3 | 8.6 | 6.42 | 1155.90 | -7.96 |
| 6 | 1105-12 | 65 | 67.0 | 3.5 | 14.5 | 7.8 | 6.30 | 1126.95 | -10.26 |
| 7 | 1110反-3 | 65 | 86.0 | 3.4 | 9.5 | 5.5 | 6.93 | 899.70 | -28.36 |
| 8 | 冀绿7号(CK) | 65 | 67.0 | 3.1 | 17.5 | 8.9 | 6.43 | 1255.80 |
“*”表示在P < 0.05水平差异显著。
“*”indicates significant difference at the P < 0.05 level.
参试品系田间枯萎病抗性调查于收获期进行，根据植株叶片黄化面积及茎基部维管束的变化评价其抗性。结果（图2）显示，6份高抗品系叶片均未出现黄化或病斑，茎基部也未出现维管束变褐，8份抗性品系的叶片出现轻度黄化或枯萎，但茎基部未出现维管束变褐，证明这些抗病品系在田间全生育期表现抗枯萎病，与苗期鉴定结果一致。
图2
图2
绿豆品系田间病圃收获期维管束及植株表型
(a) 感病品系茎基部维管束；(b) 感病品系成熟期植株表型；(c) 抗病品系茎基部维管束；(d) 抗病品系成熟期植株表型。
Fig.2
Vascular bundle and plant phenotype of mung bean line materials at harvest stage in field disease nursery
(a) Vascular bundle at stem base of susceptible line; (b) Plant phenotype of susceptible line at maturity; (c) Vascular bundle at stem base of resistant line; (d) Plant phenotype of resistant line at maturity.
2.2.2 河北省绿豆新品种区域试验
抗性品系HN0802-1-2-2-1-1-4参加2019-2020年河北省绿豆新品种区域试验。由表3可以看出，抗性品系HN0802-1-2-2-1-1-4与保绿201321-7的产量显著高于其他参试品系，其中HN0802-1-2-2-1-1-4产量最高，为1817.6 kg/hm2，较对照增产9.80%。
表3 河北省绿豆区域试验参试品种（系）产量差异多重比较结果
Table 3
| 序号 Number | 品种（系） Variety (line) | 产量 Yield (kg/hm2) | 比对照增产 Yield increase than control (%) |
|---|---|---|---|
| 1 | HN0802-1-2-2-1-1-4 | 1817.6a | 9.80 |
| 2 | 保绿201321-7 | 1765.8ab | 6.67 |
| 3 | 保绿942（CK） | 1655.4bc | 0.00 |
| 4 | 保绿201347-6 | 1631.2c | -1.46 |
| 5 | 廊绿2号 | 1632.6c | -1.38 |
| 6 | 廊绿1号 | 1621.6c | -2.04 |
| 7 | 冀绿0911-1-2 | 1611.2c | -2.67 |
| 8 | 冀绿抗1004-38-2-3-2-3 | 1569.6c | -5.18 |
| 9 | 保绿201326-1 | 1565.4c | -5.44 |
不同小写字母表示差异达显著水平（P < 0.05）。
Different lowercase letters indicate significant difference (P < 0.05).
表4 HN0802-1-2-2-1-1-4在区域试验中各试点产量结果
Table 4
| 区域 Region | 试点 Site | 2019年平均产量 Average yield in 2019 (kg/hm2) | 2019年增产率 Yield increase in 2019 (%) | 2020年平均产量 Average yield in 2020 (kg/hm2) | 2020年增产率 Yield increase in 2020 (%) | 2年平均产量 Average yield in two years (kg/hm2) | 2年平均增产率 Average yield increase in two years (%) | 位次 Ranking |
|---|---|---|---|---|---|---|---|---|
| 夏播区 Summer sowing area | 石家庄 | 2078.00 | 12.26 | 1885.70 | 2.82 | 1981.90 | 7.57 | 1 |
| 保定 | 1985.00 | 0.40 | 1787.00 | 0.51 | 1886.00 | 0.45 | 5 | |
| 唐山 | 1655.00 | -2.42 | 1769.20 | 8.35 | 1712.10 | 2.86 | 3 | |
| 廊坊 | 1157.00 | 3.58 | 1736.40 | 8.11 | 1446.70 | 6.25 | 1 | |
| 衡水 | 553.00 | -27.90 | 1750.00 | 7.51 | 1151.50 | -3.83 | 6 | |
| 邯郸 | 2740.00 | 3.01 | 1310.00 | 22.43 | 2025.00 | 8.58 | 1 | |
| 平均Average | 1694.70 | -1.85 | 1706.38 | 8.29 | 1700.53 | 3.65 | 1 | |
| 春播区 Spring sowing area | 张家口 | 1983.30 | -16.00 | 2081.80 | 37.95 | 2032.60 | 5.04 | 1 |
| 阳原 | 2690.00 | 56.40 | 1920.00 | 50.00 | 2305.00 | 53.67 | 1 | |
| 平均Average | 2336.65 | 20.20 | 2000.90 | 43.98 | 2168.80 | 29.36 | 1 |
表5 参试品种丰产性及其稳定性分析
Table 5
| 序号 Number | 品种（系） Variety (line) | 丰产性参数Yield index | 稳定性参数Stability index | 适应地区 Suitable sites | 综合评价 Comprehensive evaluation | |||
|---|---|---|---|---|---|---|---|---|
| 产量Yield (kg/hm2) | 效应 | 方差 | 变异度 | |||||
| 1 | HN0802-1-2-2-1-1-4 | 1817.6 | 0.167 | 0.024 | 8.402 | E1~E8 | 很好 | |
| 2 | 保绿201321-7 | 1765.8 | 0.119 | 0.021 | 8.082 | E1~E8 | 好 | |
| 3 | 保绿942（CK） | 1655.4 | 0.004 | 0.024 | 9.147 | E1~E8 | 一般 | |
| 4 | 保绿201347-6 | 1631.2 | -0.022 | 0.023 | 9.094 | E1~E8 | 一般 | |
| 5 | 廊绿2号 | 1632.6 | -0.023 | 0.010 | 6.138 | E1~E8 | 一般 | |
| 6 | 廊绿1号 | 1621.6 | -0.033 | 0.009 | 5.663 | E1~E8 | 一般 | |
| 7 | 冀绿0911-1-2 | 1611.2 | -0.042 | 0.061 | 15.177 | E1~E8 | 一般 | |
| 8 | 冀绿抗1004-38-2-3-2-3 | 1569.6 | -0.083 | 0.028 | 10.459 | E1~E8 | 较差 | |
| 9 | 保绿201326-1 | 1565.4 | -0.088 | 0.016 | 8.014 | E1~E8 | 较差 | |
E1~E8依次为石家庄、保定、唐山、廊坊、衡水、邯郸、张家口、阳原。
E1-E8 represent the locations of Shijiazhuang, Baoding, Tangshan, Langfang, Hengshui, Handan, Zhangjiakou and Yangyuan, respectively.
2.2.3 HN0802-1-2-2-1-1-4综合农艺性状
区域试验结果（表6）表明，HN0802-1-2-2-1-1-4为早熟种，株型直立，抗倒性强，结荚集中，成熟一致。其中，生育期春播区77.0 d，夏播区68.0 d，株高46.6~55.6 cm，主茎分枝数3.9~4.1，主茎节数9.0~9.9，单株结荚27.6~30.7，荚长11.8~12.2 cm，籽粒绿色，有光泽，饱满整齐，百粒重6.8~7.0 g。
表6 HN0802-1-2-2-1-1-4主要农艺性状
Table 6
| 区域 Region | 生育期 Growth period (d) | 株高 Plant height (cm) | 单株分枝数 Branch number per plant | 主茎节数 The number of main stem sections | 单株荚数 Pod number per plant | 荚长 Pod length (cm) | 单荚粒数 Grain number per pod | 百粒重 100-seed weight (g) |
|---|---|---|---|---|---|---|---|---|
| 春播区Spring sowing area | 77.0 | 46.6 | 4.1 | 9.9 | 30.7 | 11.8 | 9.9 | 7.0 |
| 夏播区Summer sowing area | 68.0 | 55.6 | 3.9 | 9.0 | 27.6 | 12.2 | 10.5 | 6.8 |
2.2.4 HN0802-1-2-2-1-1-4生产试验产量表现
抗性品系HN0802-1-2-2-1-1-4提交参加了2020年在石家庄、保定、唐山、廊坊、张家口5个试点进行的绿豆新品种生产试验。结果（表7）表明，该品系在5个试点均表现增产，增产率范围为2.39%~ 51.72%，5个试点平均产量为1937.10 kg/hm2，较保绿942（CK）增产23.77%。
表7 HN0802-1-2-2-1-1-4生产试验产量表现
Table 7
| 试点 Site | 产量 Yield (kg/hm2) | 对照种产量 Yield of CK (kg/hm2) | 较CK增产率 Yield increase than CK (%) |
|---|---|---|---|
| 石家庄Shijiazhuang | 2508.00 | 1653.00 | 51.72 |
| 保定Baoding | 1750.80 | 1434.90 | 22.02 |
| 唐山Tangshan | 1794.00 | 1662.00 | 7.94 |
| 廊坊Langfang | 1545.00 | 1509.00 | 2.39 |
| 张家口Zhangjiakou | 2088.00 | 1566.00 | 33.33 |
| 平均Average | 1937.10 | 1565.10 | 23.77 |
3 讨论
冀绿23是由潍绿9002-341、冀绿98-1和冀绿2号为亲本杂交选育而成。经鉴定，亲本之一潍绿9002-341病情指数为6.01，为高抗枯萎病品种，经连续鉴定选择，冀绿23很好地保留了其亲本的枯萎病抗性。因此，筛选和利用抗性种质是培育抗病品种的前提。
在抗病育种工作中，理想的抗性品种既要保证品种对枯萎病抗性的稳定性，又要提高对枯萎病的抗性级别，使两者完美的结合起来。但在现阶段抗病资源缺乏的情况下，除了利用高抗材料外，也可以对表现抗和中抗的材料加以利用。这些材料虽然不能完全抵抗病菌侵染，但是这些材料具有延迟发病和降低病害危害程度的作用，且往往具有广谱抗性和丰富的抗源，在抗病品种和育种材料匮乏的情况下，这些材料的保存和合理利用也显得尤为重要。
4 结论
利用剪根浸根接种法对81份绿豆新品系进行苗期枯萎病抗性鉴定，创制出6份高抗（HR）和8份抗性（R）种质；创制出的14份种质材料通过高级产量比较鉴定试验、省区域试验及生产试验，选育出了抗枯萎病、高产稳产、优质的绿豆新品种冀绿23。
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PMID:30732107
[本文引用: 1]
Fusarium wilt, caused by race 4 of Fusarium oxysporum f. sp. vasinfectum, is a critically important disease problem in California cotton (Pima, Gossypium barbadense; Upland, G. hirsutum). Because few cultivars with resistance to race 4 are available, alternative management strategies for this disease are needed. Four soil treatments (50:50 methyl-bromide + chloropicrin as a positive control; 60:40 chloropicrin + 1,3-dichloropropene; 6 weeks of solarization; and metam-sodium) were evaluated for efficacy against race 4 in a naturally infested, heavy clay soil. Treatments were evaluated based on plant mortality, height, number of mainstem nodes, vascular discoloration ratings, and soil counts of F. oxysporum. Two cultivars each of Pima and Upland cotton varying in resistance to race 4 were used. Plant mortality was lowest in methyl-bromide + chloropicrin, solarization, and chloropicrin + 1,3-dichloropropene treatments, and highest in the nontreated and metam-sodium treatments. Although most plant mortality occurred within 5 weeks after planting, substantial mortality of the susceptible Pima cultivar DP 744 accumulated for up to 10 weeks. Seven to eight weeks after planting, plants in methyl-bromide + chloropicrin and chloropicrin + 1,3-dichloropropene treatments were taller and had more mainstem nodes than in other treatments. Vascular discoloration was reduced in methyl-bromide + chloropicrin and solarization treatments compared with the nontreated control, metam-sodium, and chloropicrin + 1,3-dichloropropene treatments. Soil counts of F. oxysporum were significantly reduced only in the methylbromide + chloropicrin, chloropicrin + 1,3-dichloropropene, and solarization treatments. Six weeks of solarization and 60:40 chloropicrin + 1,3-dichloropropene (295 liters a.i./ha) proved effective for reducing Fusarium wilt of cotton in heavy clay soil.
Evaluation of trichoderma species against Fusarium oxysporum f. sp. ciceris for integrated management of chickpea wilt
DOI:10.1016/j.biocontrol.2006.06.006 URL [本文引用: 1]
Enhancement of biocontrol potential of Pseudomonas aeruginosa and Paecilomyces lilacinus against root rot of mung bean by a medicinal plant Launaea nudicaulis L.
Reservoir and non-reservoir hosts of bean-wilt pathogen, Fusarium oxysporum f. sp. phaseoli
Legume improvement program at AVRDC-the world vegetable center: impact and future prospects
DOI:10.5937/ratpov51-5488 URL [本文引用: 1]
普通菜豆镰孢菌枯萎病研究进展
DOI:10.13430/j.cnki.jpgr.20200930001
[本文引用: 1]
镰孢菌枯萎病是目前我国普通菜豆生产中发生的重要病害之一。本文综述了普通菜豆镰孢菌枯萎病病原菌鉴定、发病规律、防治措施、种质资源鉴定和抗性遗传研究等方面的研究进展，表明目前镰孢菌枯萎病的研究比较滞后，抗性种质资源和基因资源匮乏，特别是抗性分子机制的研究更是空白。今后应加强对现有种质资源的抗性评价，并从中鉴定优异基因或等位变异，进而解析其抗性分子机制，促进普通菜豆抗镰孢菌枯萎病抗性育种的发展。
Fusarium wilt of cotton: management strategies
DOI:10.1016/j.cropro.2015.01.014 URL [本文引用: 1]
Pathotype diversity of Fusarium oxysporum f. sp. mungcola causing wilt on mungbean (Vigna radiata)
DOI:10.1071/CP20164
URL
[本文引用: 1]
\nFusarium wilt, caused by Fusarium oxysporum f. sp. mungcola (Fom), is an increasingly serious disease of mungbean (Vigna radiata (L.) R.Wilczek) in China. Pathogenic variability has been observed among Fom isolates; however, there are no reports describing Fom races or pathotypes. Thus, this study was conducted with the aim of developing a set of pathotype differentials to reveal Fom pathotype diversity by assessing virulence variability of Fom isolates. First, 105 mungbean cultivars were screened against a standard virulent Fom isolate (F08). Eleven of the 105 cultivars were selected as candidate differentials of Fom according to resistance phenotype and genetic background. Second, the resistance of the 11 candidate differential cultivars was tested against 30 Fom isolates from different geographical origins in China. Highly significant differences were observed among isolate × cultivar interaction patterns, indicating that pathotype differentiation exists in Fom isolates. Based on the different reaction patterns combining with genetic background, seven of the 11 cultivars were selected to constitute a set of differential hosts of Fom pathotype, used to distinguish pathotypes of 84 Fom isolates from different geographical regions by evaluating the virulence reaction pattern. The results showed that the 84 Fom isolates were defined as 12 pathotypes. Finally, we tried to confirm whether the 12 Fom pathotypes could be distinguished by a PCR-based diagnostic method based on the two genes (SIX6 and SIX11) reported to be associated with Fom pathogenicity. However, the Fom pathotype could not be distinguished by variation of the PCR products or their resulting sequences of the two genes. This is the first study to develop a set of Fom pathotype differential hosts and identify 12 Fom pathotypes, which provides important information for resistance breeding and disease control.\n
Breeding for Fusarium wilt race 4 resistance in cotton under field and greenhouse conditions
A major gene for resistance to carbendazim, in field isolates of Gibberella zeae
DOI:10.1080/07060660509507194 URL [本文引用: 1]
