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植(zhi)物病原镰刀菌产(chan)生的毒素(su)种类及其危(wei)害

张嘉城 方香玲 南志标

引用本文: 张嘉城,方香玲,南志标. 植物病原镰刀菌产生的毒素种类及其危害. 欧宝体育, 2021, 38(8): 1513-1524 doi: shu
Citation:  ZHANG J C, FANG X L, NAN Z B. Types and effects of toxins produced by plant pathogenic fungi . Pratacultural Science, 2021, 38(8): 1513-1524 doi: shu

植物病原镰刀菌产生的毒素种类及其危害

    作者简介: 张嘉城(1998-),男,辽宁省瓦房店人,在读硕士生,研究方向为苜蓿根腐病。E-mail: zhangjch20@webs-seo.com
    通讯作者: 方香玲(1984-),女,河南平顶山人,教授,博士,研究方向为苜蓿根腐病。E-mail: xlf@webs-seo.com
  • 基金项目: 国家自然科学基金项目(31802126);青海省科技厅基础研究计划创新团队项目(2021-ZJ-901);国家绿肥产业技术体系(CARS-22)

摘要: 镰刀菌(Fusarium spp.)是多种重要农作物的病原体,不仅可造成农作物产量和品质的严重损失还可在离体培养条件下或植物寄主体内产生一系列被称为镰刀菌毒素的次生代谢产物。这些毒素一方面作为致病因子与镰刀菌对宿主植物的致病力密切相关,另一方面可导致家畜生产性能下降和相关病症的出现,进而影响农业生态系统并对人类健康造成威胁。鉴于镰刀菌毒素对农作物生产的影响及其对家畜和人类的毒性作用,目前已有较多欧宝体育镰刀菌侵染粮食作物后产生毒素种类的研究,但欧宝体育镰刀菌侵染豆科牧草后产生的毒素种类以及毒素在镰刀菌对豆科牧草致病力方面作用的研究则较少。本文对引起主要粮食和饲料作物病害的常见镰刀菌物种产生的主要毒素,以及这些毒素对植物、家畜和人类的危害进行了综述,并对豆科牧草中镰刀菌毒素的研究前景及意义进行了展望。

English

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    表 1  镰刀菌在人工培养条件下的代谢毒素种类

    Table 1.  Types of toxins produced by Fusarium species in vivo

    镰刀菌
    Fusarium species
    培养基类型a
    Culture medium
    毒素b
    Toxins
    层出镰刀菌 F. proliferatum 玉米 Maize FB1, MON, BEA
    黄色镰刀菌 F. culmorum 小麦,玉米 Wheat, maize DON, 3-ADON, NIV, ZEA, MON
    梨孢镰刀菌 F. poae 玉米,稻米,小米 Maize, rice, millet, PSA T-2, HT-2, NEO, DAS, NIV, FX, BEA, ENN-B
    拟枝孢镰刀菌 F. sporotrichioides 玉米,稻米 Maize, rice, PSA T-2, HT-2, T-2 triol, T-2 tetraol, NEO
    DAS, DON, ZEA, MON, BEA
    三线镰刀菌 F. tricinctum 稻米 Rice ENN-A, ENN-A1, ENN-B, ENN-B1, MON, BEA
    木贼镰刀菌 F. equiseti 玉米 Maize DON, ZEA, BEA
    轮枝样镰刀菌 F. verticillioides 玉米,稻米 Maize, rice FB1, FB2, MON, BEA, FA
    亚粘团镰刀菌 F. subglutinans 玉米,稻米 Maize, rice FB1, FB2, ZEA, MON, BEA, FA
    茄病镰刀菌 F. solani 玉米,稻米 Maize, rice, PDA FB1, FB2, MON, FA
    尖孢镰刀菌 F. oxysporum 玉米,稻米 Maize, rice DON, ENN-A, ENN-A1, ENN-B1, MON, BEA, FA
    燕麦镰刀菌 F. avenaceum 玉米,稻米 Maize, rice MON, BEA, ENN-A1, ENN-B, ENN-B1
    半裸镰刀菌 F. semitectum 玉米,稻米 Maize, rice T-2, DAS, ZEA, MON, BEA
    禾谷镰刀菌 F. graminearum 玉米 Maize DAS, DON, 3-ADON, ZEA
    藤仓镰刀菌 F. fujikuroi 玉米,稻米 Maize, rice, PDA FB1, FB2, MON, BEA, FA
    锐顶镰刀菌 F. acuminatum 稻米 Rice T-2, HT-2, NEO, MON, ENN-B
     表1根据参考文献[7-13] 汇总。a PDA: 马铃薯葡萄糖琼脂培养基;PSA: 马铃薯蔗糖琼脂培养基。b T-2,T-2毒素;HT-2,HT-2毒素;T-2 triol,T-2三醇;T-2 tetraol,T-2四醇;NEO,新茄病镰刀菌烯醇;DON,脱氧雪腐镰刀菌烯醇;3-ADON,3-乙酰脱氧雪腐镰刀菌烯醇;15-ADON,15-乙酰脱氧雪腐镰刀菌烯醇;NIV,雪腐镰刀菌烯醇;FX,镰刀菌烯酮;DAS,蛇形毒素;FB: 伏马菌素;FB1,伏马菌素B1;FB2,伏马菌素B2;MON,串珠镰刀菌素;BEA,白僵菌素;ZEA,玉米赤霉烯酮;ENN-A,恩镰孢菌素A;ENNs: 恩镰孢菌素;ENN-A1,恩镰孢菌素A1;ENN-B,恩镰孢菌素B;ENN-B1,恩镰孢菌素B1;FA,镰刀菌酸;下同。
     Table 1 was summarized based on references [7-13] . a PDA: potato dextrose agar medium; PSA: potato sucrose agar medium. b T-2: T-2 toxin; HT-2: HT-2 toxin; T-2 triol; T-2 tetraol; NEO: neosolaniol; DON: deoxynivalenol; 3-ADON: 3-acetyldeoxynivalenol; 15-ADON: 15-acetyldeoxynivalenol; NIV: nivalenol; DAS: diacetoxyscirpenol; FX: fusarenon-X; FB: fumonisins; FB1: fumonisin B1; FB2: fumonisin B2; MON: moniliformin; BEA: beauvericin; ZEN: zearalenone; ENNs: enniatins; ENN-A: enniatin A; ENN-A1: enniatin A1; ENN-B: enniatin B; ENN-B1: enniatin B1; FA: fusaric acid; this is applicable for the following tables as well.
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    表 2  镰刀菌在植物体内的代谢毒素种类

    Table 2.  Types of toxins produced by Fusarium species in planta

    镰刀菌
    Fusarium species
    寄主植物
    Host plant
    毒素
    Toxins
    层出镰刀菌 F. proliferatum 玉米 Maize FB1, FB2, BEA
    黄色镰刀菌 F. culmorum 小麦,大麦 Wheat, barley HT-2, DON, 3-ADON, NIV, ZEA, BEA, ENN-A
    ENN-A1, ENNB, ENNB1
    梨孢镰刀菌 F. poae 燕麦,大麦,玉米,小麦
    Oat, barley, maize, wheat
    HT-2, DAS, DON, NIV, BEA, ENN-A, ENN-A1
    ENN-B, ENN-B1
    拟枝孢镰刀菌 F. sporotrichioides 小麦,大麦 Wheat, barley T-2, HT-2, T-2 tetraol, NEO, DON, 3-ADON, NIV
    BEA, ENN-A1, ENN-B, ENN-B1
    三线镰刀菌 F. tricinctum 小麦,大麦 Wheat, barley DON, NIV, BEA, ENN-A, ENN-A1, ENN-B, ENN-B1
    木贼镰刀菌 F. equiseti 小麦,大麦,燕麦, 豌豆, 苜蓿
    Wheat, barley oat, pea, alfalfa
    NIV, ZEA
    轮枝样镰刀菌 F. verticillioides 玉米 Maize DON, FB1, FB2, ZEA
    亚粘团镰刀菌 F. subglutinans 玉米 Maize DON, MON, BEA, ZEA
    茄病镰刀菌 F. solani
    尖孢镰刀菌 F. oxysporum 香蕉,黄瓜 Banana, cucumber BEA, FA
    燕麦镰刀菌 F. avenaceum 小麦,大麦,小黑麦
    Wheat, barley, triticale
    HT-2, DON, NIV, MON, BEA, ENN-A, ENN-B
    ENN-B1
    半裸镰刀菌 F. semitectum
    禾谷镰刀菌 F. graminearum 小麦,大麦 Wheat, barley HT-2, DON, 3-ADON, 15-ADON, NIV, ZEA, BEA
    ENN-A1, ENN-B, ENN-B1
    藤仓镰刀菌 F. fujikuroi
    锐顶镰刀菌 F. acuminatum 小麦 Wheat T-2, HT-2
     表2根据参考文献[10-11, 14-24] 汇总。
     Table 2 was summarized based on references [10-11, 14-24].
    下载: 导出CSV

    表 3  镰刀菌毒素的植物毒性

    Table 3.  Phytotoxic effects of Fusarium toxins

    毒素
    Toxin
    植物材料
    Plant material
    浓度
    Concentration/(μg·mL−1)
    毒性效应
    Toxicity effects
    参考文献
    Reference
    DON 小麦胚芽鞘
    Wheat coleoptile
    0.30~296.00 抑制生长
    Inhibition of growth
    [62]
    3-ADON 小麦胚芽鞘
    Wheat coleoptile
    0.34~338.00 抑制生长
    Inhibition of growth
    [62]
    NIV 小麦幼苗
    Wheat seedling
    25.00~150.00 抑制生长
    Inhibition of growth
    [63]
    T-2 小麦幼苗
    Wheat seedling
    1.00 抑制生长
    Inhibition of growth
    [64]
    玉米愈伤组织
    Corn callus
    1.00~100.00 抑制生长
    Inhibition of growth
    [65]
    FB1 番茄幼苗叶片
    Tomato leaf
    0.07~72.10 叶片组织坏死
    Necrosis of leaf tissue
    [66]
    番茄幼苗
    Tomato seedling
    0.07~72.10 抑制生长
    Inhibition of growth
    [66]
    玉米幼苗
    Corn seeding
    0.07~72.10 抑制生长
    Inhibition of growth
    [66]
    黄瓜幼苗
    Cucumber seedling
    200.00 叶片黄化
    Leaf chlorosis
    [67]
    大豆幼苗
    Soybean seedling
    1000.00 叶片坏死斑
    Necrotic spot of leaf
    [67]
    棉花幼苗
    Cotton seedling
    500.00 叶片坏死斑
    Necrotic spot of leaf
    [67]
    ENN-B 小麦幼苗
    Wheat seedling
    10.00~80.00 抑制生长
    Inhibition of growth
    [68]
    ENNs 马铃薯块茎
    Potato tuber
    5.00~100.00a 块茎组织坏死
    Necrosis of tuber tissue
    [69]
    FA 香蕉幼苗
    Banana plantlet
    89.50~179.00 根茎、假茎变色,叶片萎蔫
    Rhizome and pseudostems discoloration, leaves wilt
    [70]
    棉花幼苗
    Cotton seedling
    89.50~1432.00 叶片坏死斑
    Leaf necrosis
    [71]
    黄瓜幼苗
    Cucumber seedling
    100.00 幼苗萎蔫
    Seedlings wilt
    [72]
    番茄幼苗
    Tomato seedling
    89.50~179.00 幼苗萎蔫
    Seedlings wilt
    [60]
    ZEA 玉米胚
    Corn embryo
    5.00 抑制生长
    Inhibition of growth
    [73]
    MON 玉米愈伤组织
    Corn callus
    10.00~100.00 抑制生长
    Inhibition of growth
    [65]
     a 施加在每片马铃薯块茎的毒素浓度单位: μg。
     a The concentration unit of the toxin applied to each potato slice: μg.
    下载: 导出CSV

    表 4  家畜摄入镰刀菌毒素后的症状

    Table 4.  Livestock symptoms after Fusarium toxin intake

    毒素
    Toxin
    家畜
    Livestock
    毒素来源
    Toxin source
    剂量
    Dosage/(mg·kg−1)
    时间
    Time/d
    症状
    Symptoms
    参考文献
    Reference
    T-2
    Swine
    发霉玉米
    Moldy corn
    3.20 21 摄食减少
    Reduced feed intake
    [75]
    纯化毒素
    Purified toxin
    > 4.00 14 皮炎
    Dermatitis
    [75]
    纯化毒素
    Purified toxin
    > 2.00 21 增重减少
    Reduced weight gain
    [75]
    纯化毒素
    Purified toxin
    > 2.00 21 摄食减少
    Reduced feed intake
    [75]
    绵羊
    Lamb
    纯化毒素
    Purified toxin
    0.60a 21 腹泻
    Diarrhea
    [76]

    Cattle
    0.64 20 血便、肠炎、皱胃和瘤胃溃疡
    Bloody feces, enteritis, abomasal and ruminal ulcers
    [76]
    NIV 肉鸡
    Broiler
    纯化毒素
    Purified toxin
    > 6.00 20 摄食和增重减少
    Reduced feed intake and weight gain
    [77]
    DON
    Swine
    发霉玉米
    Moldy corn
    > 1.34 21 摄食和增重显著减少
    Reduced feed intake and weight gain
    [78]
    发霉玉米
    Moldy corn
    > 11.90 21 完全拒食
    Complete feed refusal
    [78]
    发霉玉米
    Moldy corn
    > 19.70 21 呕吐
    Vomiting
    [78]
    FB1
    Swine
    发霉玉米
    Moldy corn
    166.00 6 急性肺水肿
    Pulmonary edema
    [79]
    肉鸡
    Broiler
    轮枝样镰刀菌
    F. verticillioides
    > 100.00 21 腹泻和增重减少
    Diarrhea and reduced weight gain
    [79]
    纯化毒素
    Purified toxin
    125.00 14 死亡
    Death
    [80]
    FB 绵羊
    Lamb
    轮枝样镰刀菌
    F. verticillioides
    45.50a 7 死亡
    Death
    [79]
    ZEA
    Swine
    发霉玉米
    Moldy corn
    > 3.00 14 高雌性素症
    Hyperestrogenism
    [81]
    纯化毒素
    Purified toxin
    > 60.00 42 不孕
    Infertility
    [82]
    奶牛
    Dairy
    纯化毒素
    Purified toxin
    250 mgb 产奶量减少,不孕
    Reduced milk production and infertility
    [81]
    绵羊
    Lamb
    纯化毒素
    Purified toxin
    12.50 10 排卵率和生育力下降
    Reduced ovulation rate and fertility
    [83]
    火鸡
    Turkey
    纯化毒素
    Purified toxin
    100.00 21 产蛋量减少
    Reduced egg production
    [81]
    MON 肉鸡
    Turkey
    纯化毒素
    Purified toxin
    27.00 7 增重减少和死亡
    Reduced weight gain and death
    [80]
    火鸡
    Turkey
    藤仓镰刀菌
    F. fujikuroi
    100.00 21 摄食量和增重减少
    Reduced feed intake and weight gain
    [84]
    母鸡
    Hen
    藤仓镰刀菌
    F. fujikuroi
    100.00 28 产蛋量和增重减少
    Reduced egg production and weight gain
    [85]
    FA
    Swine
    纯化毒素
    Purified toxin
    200 mgb 呕吐
    Vomiting
    [86]
     a 摄入毒素剂量单位(体重基础上): mg·kg−1b 毒素通过明胶胶囊一次性饲喂家畜。
      a Intake dosage unit (body weight basis): mg·kg−1. b Toxins are fed to livestock at one time via gelatin capsules.
    下载: 导出CSV
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                • 通讯作者:  方香玲, xlf@webs-seo.com
                • 收稿日期:  2020-12-30
                • 接受日期:  2021-04-06
                • 网络出版日期:  2021-07-06
                • 刊出日期:  2021-08-15
                通(tong)讯作者: 陈斌, bchen63@163.com
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