▷ 苹果轮纹病病害名称之规范

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苹果轮纹病(apple ring rot)严重危害果实和枝干，导致苹果果实腐烂、枝干表皮疣突、溃疡，危害幼树导致整树枯死，与苹果树腐烂病和早期落叶病被并称为我国苹果产业的三大病害(国立耘等 2009;李保华等 2013)。

该病于1929年在辽宁发现(景学富等 1963)。目前在我国大部分苹果产区普遍发生，尤其在渤海湾苹果产区的河北、山东、辽宁等省和黄河故道区河南和江苏省等地严重发生(胡清玉等 2016)。2001年烟台地区苹果轮纹病大发生，一般果园烂果率在30%-50%，部分果园甚至绝收(董娟华和李保华 2009)。随着苹果套袋技术的推广，果实发病得到有效控制，但用药的减少导致枝干发病愈趋严重(Tang et al. 2012)。近些年，随着气候、栽培制度及大量异地苗木调运等原因，该病在黄土高原区的陕西、山西、甘肃省矮化果园危害也在加重，幼树大面积死亡现象常有发生。

对于“苹果轮纹病”是一种病害还是几种病害，国内外一直缺乏一致的看法。在我国相关病害名称的使用十分混乱，如苹果轮纹病、苹果干腐病、苹果果实轮纹病、苹果枝干轮纹病及轮纹烂果病等不同名称被广泛使用。近些年来，随着多基因系统发育分析及全基因组分析技术在该类群研究中的应用，明确了苹果轮纹病是一种复合病害。本文根据近些年来国内外研究新进展，对病害的汉语学称、英文名称、病原的拉丁学名等提出了规范用法。

1 相关病原的鉴定与分类研究

1.1 苹果病原葡萄座腔菌分类鉴定

葡萄座腔菌属Botryosphaeria Ces. et de Not.由de Notaris (1863)建立，其中包含了葡萄座腔菌B. dothidea，其最早发现于蔷薇属Rosa植物，但未指定属的模式。de Notaris (1863)增加更多种到Botryosphaeria，其中包括贝林格葡萄座腔菌B. berengeriana。von Arx & Muller (1954)整理Botryosphaeria属的种，认为茶藨子葡萄座腔菌B. ribis是B. dothidea的异名。Barr (1972)指定B. dothidea为属的候选模式(lectotype)。近些年来，随着分子系统学研究技术的发展，B. dothidea被广泛认可，B. berengeriana属于其异名(Smith et al. 2001;Slippers et al. 2004, 2014;Phillips et al. 2013)。

对于苹果相关的病原，Grossenbacher & Dugger (1911)将引起茶藨子溃疡的病原定名为B. ribis。Putterill (1919)在南非发现苹果干腐病，将病原鉴定为B. mali (无性阶段为有子座的Dothiorella mali E.和无子座的Macrophoma spp.)。Fenner (1925)和Shear et al. (1925)在美国报道苹果果实白腐病(white rot, Botryosphaeria rot or bot rot)，病原为B. ribis，并认为Dothiorella mali与B. ribis是同一个种。von Arx & Muller (1954)曾提出B. ribis为B. dothidea异名，因而B. dothidea被欧美学者作为苹果白腐病的病原广泛采纳(Sutton et al. 2014)。但是，von Arx & Muller (1975)又提出B. ribis是B. berengeriana的异名，因此，小金泽硕城和佐久间勉(1984)将日本干腐型病菌鉴定为B. berengeriana。该名称被日本和部分中国学者采用(陈策 1999)。Xu et al. (2015)采用多基因核苷酸系统发育明确B. dothidea为我国苹果轮纹病主要病原。目前大量研究显示，B. ribis为独立种，但被归属于新壳梭孢属Neofusicoccum;Dothiorella不是葡萄座腔菌属的无性阶段，而是一个独立的属;Fusicoccum aesculi被认为是B. dothidea的无性阶段名称(Zhang W et al. 2021;Zhang Y et al. 2021)，根据“One Fungus One Name”新法规，该名称不再作为正式名称。葡萄座腔菌B. dothidea是我国苹果轮纹病主要病原之一，其汉语名称、拉丁学名及异名如下：

Botryosphaeria dothidea (Moug.) Ces. & De Not. 葡萄座腔菌

≡ Sphaeria dothidea Moug.

= Botryosphaeria berengeriana De Not.

= Fusicoccum aesculi Corda

1.2 苹果病原锹冢葡萄座腔菌分类鉴定

轮纹病(ring rot)于1907年最早于日本在梨果实上发现。1921年，锹冢在苹果枝条上报道由大茎点霉Macrophoma引起的苹果疣皮病，1930年原摄祐将疣皮病病原定名为锹冢大茎点霉M. kuwatsukai Hara (北岛博 1989)，该菌可以侵染苹果果实引起轮纹症状，因而将苹果疣皮病、瘤状粗皮病与引起的果实腐烂病害作为一种病害——轮纹病。Nose (1933)在梨上发现M. kuwatsukai的有性时期，命名为梨生囊孢Physalospora piricola Nose，后被修订为梨生球座菌Guignardia pyricola (Yamamoto 1961)。小金泽硕城和佐久间勉(1980, 1984)在研究苹果果实腐烂病原时，认为轮纹病的病原不属于Physalospora，而属于Botryosphaeria;他们发现果实腐烂的病原除了“苹果干腐病菌B. berengeriana”外，还存在一种新病原，其形态与B. berengeriana无明显差异，但主要引起疣皮症状，定名为贝林格葡萄座腔菌梨生专化型B. berengeriana f. sp. piricola。该病原名称被日本学者广泛采用，但由于B. berengeriana被认为是B. dothidea的异名(Barr 1972;Slippers et al. 2004, 2014)，因而其种下的专化型B. berengeriana f. sp. piricola一直未被欧美和韩国学者认同。尽管如此Sutton et al. (2014)仍在其专著Compendium of apple and pear diseases and pests中将“apple ring rot”列为一个条目，对B. berengeriana f. sp. piricola进行了讨论。Xu et al. (2015)采用多基因核苷酸系统发育揭示B. berengeriana f. sp. piricola与B. dothidea存在显著的遗传分化，结合致病性、寄主范围、生物特性等多项性状，明确其为一个独立物种，并将B. berengeriana f. sp. piricola修订新组合为B. kuwatsukai。Wang et al. (2018，2021)通过全基因组序列分析，认为B. dothidea与B. kuwatsukai在大约250万年前分化为不同的种类。Zhang W et al. (2021)和Zhang Y et al. (2021)基于多基因系统发育分析和形态学分析支持B. kuwatsukai为独立物种，并认为Zhou et al. (2017)所建立的新种Botryosphaeria rosaceae Y.P. Zhou & Y. Zhang ter.为其异名。根据这些研究结果，锹冢葡萄座腔菌Botryosphaeria kuwatsukai也是我国苹果轮纹病主要病原之一，其相关异名如下：

Botryosphaeria kuwatsukai (Hara) G.Y. Sun and E. Tanaka 锹冢葡萄座腔菌

≡Macrophoma kuwatsukai Hara

= Physalospora pyricola Nose

= Guignardia pyricola (Nose) W. Yamamoto

= Botryosphaeria berengeriana De Notaris f. sp. pyricola Koganezawa & Sakuma

= Botryosphaeria rosaceae Y.P. Zhou & Y. Zhang ter.

1.3 葡萄座腔菌与锹冢葡萄座腔菌的致病性差异

葡萄座腔菌和锹冢葡萄座腔菌均可以侵染苹果枝干和果实，但是在不同的文献中，对症状描述存在明显的差异。小金泽硕城和佐久间勉(1984)接种实验显示B. berengeriana (=B. dothidea)从伤口侵入枝条引起溃疡病斑，分生孢子无伤接种不发病，而B. berengeriana f. sp. pyricola (=B. kuwatsukai)无伤分生孢子接种可以在皮孔周围形成疣状突起。陈策(1993)认为“溃疡病菌”可以从伤口和皮孔侵入，伤口侵入发病较快，皮孔侵入发病较慢;“轮纹病菌”以皮孔侵入为主，以皮孔为中心疣状隆起，后组织坏死，表现为指甲大小溃疡斑。Xu et al. (2015)采用菌饼无伤接种显示B. dothidea 产生直径为0.7-1.0 mm的小瘤，而B. kuwatsukai产生直径为3.0-4.1 mm的大瘤。Tang et al. (2012)采用来自溃疡和瘤突的不同“B. dothidea”分离物接种，也表现出大小不同的瘤突症状。一般认为干旱胁迫有利于B. dothidea引起的干腐症状的发生，最近Dong et al. (2021)用来自“瘤突的B. dothidea菌”接种，在水分胁迫条件下亦生产了溃疡症状，可能意味着B. kuwatsukai也有可能引起干腐症状。大量研究都表明葡萄座腔菌B. dothidea和锹冢葡萄座腔菌B. kuwatsukai均可以引起果实的腐烂症状，在一些苹果上表现为轮纹症状，但是两种病原引起的症状没有差异(小金泽硕城和佐久间勉1984;陈策1993;Xu et al. 2015)。

2 我国苹果轮纹病病害名称建议

苹果轮纹病可以侵染果实和枝干，引起的症状也有很大变化，病原复杂多样，病原菌的分类系统也在一直变化，导致该病害的鉴定非常困难。在我国，对该类病害的定义存在异议，导致在不同专著中的描述差异很大。一些著作作为一种病害，称为苹果轮纹病或苹果干腐病(冷怀琼1987;方中达1996;Tang et al. 2012;侯明生和黄俊斌2014;国立耘等2020)，或苹果和梨轮纹病(高学文和陈孝仁2018);一些著作作为2种病害：苹果轮纹病和苹果干腐病(浙江农业大学等1978;中国农作物病虫害编委会1979;中国农业科学院果树研究所1994;刘克均1996;张志铭1996;中国农业科学院植物保护研究所1996;王江柱等2018);还有一些描述为3种病害，分别为苹果轮纹病、苹果干腐病和果实轮纹病(或轮纹烂果病) (江苏农科院植物保护系1997;王江柱和解金斗2019;鲁传涛2021)。对病原学名的使用也较混乱，如苹果干腐病的病原有贝林格葡萄座腔菌B. berengeriana、贝克葡萄座腔菌B. bakeri、葡萄座腔菌B. dothidea及茶藨子葡萄座腔菌B. ribis等，苹果轮纹病的病原有贝林格葡萄座腔菌梨生专化型B. berengeriana f. sp. piricola、葡萄座腔菌B. dothidea、轮纹大茎点M. kuwatsukai及梨生囊壳孢Physalospora piricola等(表1)。

Table 1

表1

我国苹果轮纹病病害名称及病原拉丁学名使用情况

Table 1 Disease names of apple ring rot and Latin scientific names of pathogens used in China

著作名称Literature作者Author病害名称Disease name拉丁学名和汉语学名Scientific name and Chinese name of the pathogen《果树病理学》(1978)Fruit tree pathology (1978)浙江农业大学等Zhejiang Agricultural University, etc.苹果干腐病Apple Bot canker茶藨子葡萄座腔菌Botryosphaeria ribis苹果轮纹病Apple ring rot有性时期：梨生囊壳孢Teleomorph: Physalospora piricola无性时期：轮纹大茎点Anamorph: Macrophoma kuwatsukai《中国农作物病虫害(下册)》(1979)Crop pests in China (1979)《中国农作物病虫害》编辑委员会Chinese Editorial Committee on Crop Pests and Diseases苹果干腐病Apple Bot canker茶藨子葡萄座腔菌Botryosphaeria ribis苹果轮纹病Apple ring rot有性时期：梨生囊壳孢Teleomorph: Physalospora piricola无性时期：轮纹大茎点Anamorph: Macrophoma kuwatsukai《果树病害》(1987)Fruit tree diseases (1987)冷怀琼Leng Huaiqiong苹果轮纹病Apple ring rot有性时期：梨生囊壳孢Teleomorph: Physalospora piricola无性时期：轮纹大茎点Anamorph: Macrophoma kuwatsukai《中国果树病虫志》(1994)Compendium of fruit tree pests in China (1994)中国农业科学院果树研究所Institute of Fruit Science, Chinese Academy of Agricultural Sciences苹果干腐病Apple Bot canker大茎点型Macrophoma type小穴壳型Dothiorella type苹果轮纹病Apple ring rot轮纹大茎点Macrophoma kuwatsukai《中国农业植物病害》(1996)Agricultural plant diseases in China (1996)方中达Fang Zhongda苹果干腐病Apple Bot canker葡萄座腔菌Botryosphaeria dothidea《中国农作物病虫害(下册)》(1996)Crop pests in China (1996)中国农业科学院植物保护研究所Institute of Plant Protection, Chinese Academy of Agricultural Sciences苹果干腐病Apple Bot canker葡萄座腔菌属Botryosphaeria berengeriana苹果轮纹病Apple ring rot轮纹大茎点Macrophoma kuwatsukai著作名称Literature作者Author病害名称Disease name拉丁学名和汉语学名Scientific name and Chinese name of the pathogen刘克均Liu Kejun仁果轮纹病Pomaceous ring rot有性时期：贝氏葡萄座腔菌梨专化型Teleomorph: Botryosphaeria berengeriana f. sp. piricola无性时期：轮纹大茎点菌Anamorph: Macrophoma kuwatsukai《植物病害诊断》(1997)Plant disease diagnosis (1997)江苏农学院植物保护系Plant Protection Department, Jiangsu Agricultural College枝干干腐病Trunk Bot canker有性时期：茶藨子葡萄座腔菌Teleomorph: Botryosphaeria ribis无性时期：大茎点型Anamorph: Macrophoma type枝干轮纹病Trunk ring rot有性时期：梨生囊壳孢Teleomorph: Physalospora piricola无性时期：轮纹大茎点Anamorph: Macrophoma kuwatsukai果实轮纹病Fruit ring rot有性时期：梨生囊壳孢Teleomorph: Physalospora piricola无性时期：轮纹大茎点Anamorph: Macrophoma kuwatsukai《农业植物病理学》(2014)Agricultural plant pathology (2014)侯明生、黄俊斌Hou Mingsheng, Huang Junbin苹果轮纹病Apple ring rot有性时期：贝伦格葡萄座腔菌梨生专化型Teleomorph: Physalospora piricola无性时期：小穴壳菌Anamorph: Dothiorella gregaria《现代落叶果树病虫害诊断与防控原色图鉴》(2018)Illustrated handbook of diagnosis and prevention of pests and diseases on deciduous fruit tree (2018)王江柱、王勤英、仇贵生Wang Jiangzhu, Wang Qinying, Qiu Guisheng苹果干腐病Apple Bot canker葡萄座腔菌B. dothidea枝干轮纹病Trunk ring rot有性时期：梨生囊壳孢Teleomorph: Physalospora piricola无性时期：轮纹大茎点Anamorph: Macrophoma kuwatsukai《农业植物病理学》(2018)Agricultural plant pathology (2018)高学文、陈孝仁Gao Xuewen, Chen Xiaoren苹果和梨轮纹病Apple and pear ring rot有性时期：茶藨子葡萄座腔菌Teleomorph: Botryosphaeria dothidea无性态：七叶树壳梭孢Anamorph: Fusicoccum aesculi《苹果高效栽培与病虫害看图防治》(2019)High efficiency cultivation, disease and pest control of apple (2019)王江柱、解金斗Wang Jiangzhu, Xie Jindou干腐病Bot canker葡萄座腔菌Botryosphaeria dothidea枝干轮纹病Trunk ring rot梨生囊壳孢Physalospora piricola轮纹烂果病Ring rot梨生囊壳孢 Physalospora piricola葡萄座腔菌 Botryosphaeria dothidea《园艺植物病理学》(2022)Horticultural plant pathology (2022)国立耘、刘凤权、黄丽丽Guo Liyun, Liu Fengquan, Huang Lili苹果、梨轮纹病(苹果干腐病)Apple and pear ring rot (Bot canker)葡萄座腔菌Botryosphaeria dothidea《果树病虫诊断与防治彩色图解》(2021)Color diagram of the diagnosis and control of fruit tree diseases and pests (2021)鲁传涛Lu Chuantao苹果干腐病Apple Bot canker贝氏葡萄座腔菌Botryosphaeria berengeriana苹果轮纹病Apple ring rot轮纹大茎点菌Macrophoma kuwatsukai

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在中国和日本，B. dothidea和B. kuwatsukai两种病原均普遍存在，Xu et al. (2015)研究显示，在美国也存在两种病原菌。两种病菌侵染苹果果实，都表现腐烂病症状，没有明显差别;在枝干上B. dothidea可以表现出溃疡症状，B. kuwatsukai表现以瘤突为主，但症状受环境条件影响较大，症状常有交叉表现，因此在田间自然条件下，根据症状准确区别是哪种病原引起较困难。本研究室从溃疡或瘤突样本上均可以分离出2种病原菌(未发表)，显示自然条件下复合侵染现象普遍发生。一种病原侵染寄主不同组织器官表现不同症状，在植物病害中非常普遍，通常作为一种病害。随着对植物病害病原研究的不断深入，一种病害由多种病原引起的现象也非常普遍。如小麦赤霉病病原有燕麦镰孢Fusarium avenaceum (Fr.) Sacc.、禾谷镰孢F. graminearum Schwabe等27种镰孢菌(陆维忠等 2001)。已发现10种刺盘孢属真菌种可以引起苹果炭疽叶枯病(Velho et al. 2019;戴蓬博等 2021)。苹果煤污病(sooty blotch and flyspeck complex)的病原多达100余种(Gleason et al. 2019)。目前在欧美国家，将该类病害也作为一种“white rot (白腐病)”(Sutton et al. 2014)。基于这些原因，我们建议该类病害作为一个多病原的复合病害，病害汉语名称统称苹果轮纹病，英文使用apple ring rot。

3 存在问题与研究展望

葡萄座腔菌和锹冢葡萄座腔菌寄主范围差异很大。葡萄座腔菌寄主范围很宽，寄主植物多达280余种，引起杨树溃疡病、枣树干腐病、梨树轮纹病、猕猴桃果腐病及李树干腐病等，相关寄主在田间都可能是苹果轮纹病的初侵染来源，增加了其防治的难度(Marsberg et al. 2017;Wang et al. 2018)。锹冢葡萄座腔菌寄主范围较窄，目前已知仅可侵害苹果、梨等少数寄主，如引起梨树轮纹病(Gu et al. 2018;Xiao et al. 2022)。建议在对苹果轮纹病发生规律、防治药剂筛选及抗病育种研究中，关注2种病原菌致病性的差异、对不同杀菌剂的敏感性差异等。B. dothidea可以引起溃疡症状、瘤状突起等不同症状，这些差异是否与接种体类型、接种方法及接种的品种等相关，B. kuwatsukai通常引起瘤突症状，但在干旱胁迫下是否也可以导致枝干溃疡或干腐症状的产生等等有待进一步研究。抗病育种为防治多年生果树病害的重要途径，由于两种病原菌在苹果上都普遍发生，因此在抗病育种的靶标菌选择中，对葡萄座腔菌和锹冢葡萄座腔菌都要充分重视。

由于B. dothidea和B. kuwatsukai的ITS、ACT、tubulin等片段的核苷酸差异很小，对2个种的区别力不足，因此对病原菌的多基因系统发育分析与鉴定中，需要选择信息化差异基因，如tef、His等。Wang et al. (2018)测序并比较分析了B. dothidea和B. kuwatsukai全基因组序列，发现后者丢失了一系列次级代谢合成关键酶、碳水化合物活性酶和植物细胞壁降解酶等致病相关基因，为相关差异致病基因的功能研究提供一些指向，建议加强相关差异致病功能基因及致病机制的解析。

苹果轮纹病引起疣皮症状，大量疣状突起联合、树皮开裂最终表现“粗皮”症状。苹果树锰过剩也可以引起粗皮病，称为苹果粗皮病(internal bark necrosis, IBN)或苹果生理粗皮病。苹果生理粗皮病多数在枝条的光滑表皮上形成疹状突起，少量也在皮孔部位成疹状突起。粗皮病果园枝条韧皮部锰含量均超过200 mg/kg，个别达到600 mg/kg以上(徐圣友等 2008)。生理粗皮病在我国多发生于沿海地区，山东胶东半岛发生较为严重，该地区病原性粗皮也普遍发生。在苹果粗皮病防治过程中，需要明确是锰毒害引起的生理性病害还是苹果轮纹病引起的病原性病害。锰过多是否会加重苹果轮纹病的发生也值得进一步深入研究。

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Multi-gene genealogies and morphological data support Diplodia cupressi sp. nov., previously recognized as D. pinea f. sp. cupressi, as a distinct species

2006

Preliminary studies on the Dothideales in temperate North America

1972

... 葡萄座腔菌属Botryosphaeria Ces. et de Not.由de Notaris (1863)建立，其中包含了葡萄座腔菌B. dothidea，其最早发现于蔷薇属Rosa植物，但未指定属的模式.de Notaris (1863)增加更多种到Botryosphaeria，其中包括贝林格葡萄座腔菌B. berengeriana.von Arx & Muller (1954)整理Botryosphaeria属的种，认为茶藨子葡萄座腔菌B. ribis是B. dothidea的异名.Barr (1972)指定B. dothidea为属的候选模式(lectotype).近些年来，随着分子系统学研究技术的发展，B. dothidea被广泛认可，B. berengeriana属于其异名(Smith et al. 2001;Slippers et al. 2004, 2014;Phillips et al. 2013). ...

... 轮纹病(ring rot)于1907年最早于日本在梨果实上发现.1921年，锹冢在苹果枝条上报道由大茎点霉Macrophoma引起的苹果疣皮病，1930年原摄祐将疣皮病病原定名为锹冢大茎点霉M. kuwatsukai Hara (北岛博 1989)，该菌可以侵染苹果果实引起轮纹症状，因而将苹果疣皮病、瘤状粗皮病与引起的果实腐烂病害作为一种病害——轮纹病.Nose (1933)在梨上发现M. kuwatsukai的有性时期，命名为梨生囊孢Physalospora piricola Nose，后被修订为梨生球座菌Guignardia pyricola (Yamamoto 1961).小金泽硕城和佐久间勉(1980, 1984)在研究苹果果实腐烂病原时，认为轮纹病的病原不属于Physalospora，而属于Botryosphaeria;他们发现果实腐烂的病原除了“苹果干腐病菌B. berengeriana”外，还存在一种新病原，其形态与B. berengeriana无明显差异，但主要引起疣皮症状，定名为贝林格葡萄座腔菌梨生专化型B. berengeriana f. sp. piricola.该病原名称被日本学者广泛采用，但由于B. berengeriana被认为是B. dothidea的异名(Barr 1972;Slippers et al. 2004, 2014)，因而其种下的专化型B. berengeriana f. sp. piricola一直未被欧美和韩国学者认同.尽管如此Sutton et al. (2014)仍在其专著Compendium of apple and pear diseases and pests中将“apple ring rot”列为一个条目，对B. berengeriana f. sp. piricola进行了讨论.Xu et al. (2015)采用多基因核苷酸系统发育揭示B. berengeriana f. sp. piricola与B. dothidea存在显著的遗传分化，结合致病性、寄主范围、生物特性等多项性状，明确其为一个独立物种，并将B. berengeriana f. sp. piricola修订新组合为B. kuwatsukai.Wang et al. (2018，2021)通过全基因组序列分析，认为B. dothidea与B. kuwatsukai在大约250万年前分化为不同的种类.Zhang W et al. (2021)和Zhang Y et al. (2021)基于多基因系统发育分析和形态学分析支持B. kuwatsukai为独立物种，并认为Zhou et al. (2017)所建立的新种Botryosphaeria rosaceae Y.P. Zhou & Y. Zhang ter.为其异名.根据这些研究结果，锹冢葡萄座腔菌Botryosphaeria kuwatsukai也是我国苹果轮纹病主要病原之一，其相关异名如下： ...

Botryosphaeria diseases of apple and peach in the southeastern United States

1986

Occurrence and control of canker rot and ring rot of apple

1993

Advance in the research of apple fruit ring rot

1999

Crop pests in China

A checklist of pathogenic fungi on apple in China

2021

1863

... 植物，但未指定属的模式.de Notaris (1863)增加更多种到Botryosphaeria，其中包括贝林格葡萄座腔菌B. berengeriana.von Arx & Muller (1954)整理Botryosphaeria属的种，认为茶藨子葡萄座腔菌B. ribis是B. dothidea的异名.Barr (1972)指定B. dothidea为属的候选模式(lectotype).近些年来，随着分子系统学研究技术的发展，B. dothidea被广泛认可，B. berengeriana属于其异名(Smith et al. 2001;Slippers et al. 2004, 2014;Phillips et al. 2013). ...

Research progress of apple ring rot disease

2009

Progression of symptoms caused by Botryosphaeria dothidea on apple branches

2021

... 葡萄座腔菌和锹冢葡萄座腔菌均可以侵染苹果枝干和果实，但是在不同的文献中，对症状描述存在明显的差异.小金泽硕城和佐久间勉(1984)接种实验显示B. berengeriana (=B. dothidea)从伤口侵入枝条引起溃疡病斑，分生孢子无伤接种不发病，而B. berengeriana f. sp. pyricola (=B. kuwatsukai)无伤分生孢子接种可以在皮孔周围形成疣状突起.陈策(1993)认为“溃疡病菌”可以从伤口和皮孔侵入，伤口侵入发病较快，皮孔侵入发病较慢;“轮纹病菌”以皮孔侵入为主，以皮孔为中心疣状隆起，后组织坏死，表现为指甲大小溃疡斑.Xu et al. (2015)采用菌饼无伤接种显示B. dothidea 产生直径为0.7-1.0 mm的小瘤，而B. kuwatsukai产生直径为3.0-4.1 mm的大瘤.Tang et al. (2012)采用来自溃疡和瘤突的不同“B. dothidea”分离物接种，也表现出大小不同的瘤突症状.一般认为干旱胁迫有利于B. dothidea引起的干腐症状的发生，最近Dong et al. (2021)用来自“瘤突的B. dothidea菌”接种，在水分胁迫条件下亦生产了溃疡症状，可能意味着B. kuwatsukai也有可能引起干腐症状.大量研究都表明葡萄座腔菌B. dothidea和锹冢葡萄座腔菌B. kuwatsukai均可以引起果实的腐烂症状，在一些苹果上表现为轮纹症状，但是两种病原引起的症状没有差异(小金泽硕城和佐久间勉1984;陈策1993;Xu et al. 2015). ...

Agricultural plant diseases in China

1996

A rot of apples caused by Botryosphaeria ribi

1925

... 对于苹果相关的病原，Grossenbacher & Dugger (1911)将引起茶藨子溃疡的病原定名为B. ribis.Putterill (1919)在南非发现苹果干腐病，将病原鉴定为B. mali (无性阶段为有子座的Dothiorella mali E.和无子座的Macrophoma spp.).Fenner (1925)和Shear et al. (1925)在美国报道苹果果实白腐病(white rot, Botryosphaeria rot or bot rot)，病原为B. ribis，并认为Dothiorella mali与B. ribis是同一个种.von Arx & Muller (1954)曾提出B. ribis为B. dothidea异名，因而B. dothidea被欧美学者作为苹果白腐病的病原广泛采纳(Sutton et al. 2014).但是，von Arx & Muller (1975)又提出B. ribis是B. berengeriana的异名，因此，小金泽硕城和佐久间勉(1984)将日本干腐型病菌鉴定为B. berengeriana.该名称被日本和部分中国学者采用(陈策 1999).Xu et al. (2015)采用多基因核苷酸系统发育明确B. dothidea为我国苹果轮纹病主要病原.目前大量研究显示，B. ribis为独立种，但被归属于新壳梭孢属Neofusicoccum;Dothiorella不是葡萄座腔菌属的无性阶段，而是一个独立的属;Fusicoccum aesculi被认为是B. dothidea的无性阶段名称(Zhang W et al. 2021;Zhang Y et al. 2021)，根据“One Fungus One Name”新法规，该名称不再作为正式名称.葡萄座腔菌B. dothidea是我国苹果轮纹病主要病原之一，其汉语名称、拉丁学名及异名如下： ...

2018

Stealth pathogens: the sooty blotch and flyspeck fungal complex

2019

... 在中国和日本，B. dothidea和B. kuwatsukai两种病原均普遍存在，Xu et al. (2015)研究显示，在美国也存在两种病原菌.两种病菌侵染苹果果实，都表现腐烂病症状，没有明显差别;在枝干上B. dothidea可以表现出溃疡症状，B. kuwatsukai表现以瘤突为主，但症状受环境条件影响较大，症状常有交叉表现，因此在田间自然条件下，根据症状准确区别是哪种病原引起较困难.本研究室从溃疡或瘤突样本上均可以分离出2种病原菌(未发表)，显示自然条件下复合侵染现象普遍发生.一种病原侵染寄主不同组织器官表现不同症状，在植物病害中非常普遍，通常作为一种病害.随着对植物病害病原研究的不断深入，一种病害由多种病原引起的现象也非常普遍.如小麦赤霉病病原有燕麦镰孢Fusarium avenaceum (Fr.) Sacc.、禾谷镰孢F. graminearum Schwabe等27种镰孢菌(陆维忠等 2001).已发现10种刺盘孢属真菌种可以引起苹果炭疽叶枯病(Velho et al. 2019;戴蓬博等 2021).苹果煤污病(sooty blotch and flyspeck complex)的病原多达100余种(Gleason et al. 2019).目前在欧美国家，将该类病害也作为一种“white rot (白腐病)”(Sutton et al. 2014).基于这些原因，我们建议该类病害作为一个多病原的复合病害，病害汉语名称统称苹果轮纹病，英文使用apple ring rot. ...

A contribution to the life-history, parasitism, and biology of Botryosphaeria ribis

1911

ATMT transformation efficiencies with native promoters in Botryosphaeria kuwatsukai causing ring rot disease in pear

2018

... 葡萄座腔菌和锹冢葡萄座腔菌寄主范围差异很大.葡萄座腔菌寄主范围很宽，寄主植物多达280余种，引起杨树溃疡病、枣树干腐病、梨树轮纹病、猕猴桃果腐病及李树干腐病等，相关寄主在田间都可能是苹果轮纹病的初侵染来源，增加了其防治的难度(Marsberg et al. 2017;Wang et al. 2018).锹冢葡萄座腔菌寄主范围较窄，目前已知仅可侵害苹果、梨等少数寄主，如引起梨树轮纹病(Gu et al. 2018;Xiao et al. 2022).建议在对苹果轮纹病发生规律、防治药剂筛选及抗病育种研究中，关注2种病原菌致病性的差异、对不同杀菌剂的敏感性差异等.B. dothidea可以引起溃疡症状、瘤状突起等不同症状，这些差异是否与接种体类型、接种方法及接种的品种等相关，B. kuwatsukai通常引起瘤突症状，但在干旱胁迫下是否也可以导致枝干溃疡或干腐症状的产生等等有待进一步研究.抗病育种为防治多年生果树病害的重要途径，由于两种病原菌在苹果上都普遍发生，因此在抗病育种的靶标菌选择中，对葡萄座腔菌和锹冢葡萄座腔菌都要充分重视. ...

Investigation on the occurrence and chemical control of Botryosphaeria canker of apple in China

2009

2020

Agricultural plant pathology

2014

Survey on the occurrence and distribution of apple diseases in China

2016

Studies on the occurrence and control of trunk rot of apple tree

1963

Fruit tree diseases

1987

Research progress in apple diseases and problems in the disease management in China

2013

Pome fruits ring rot

1996

Color diagram of the diagnosis and control of fruit tree diseases and pests

2021

Study on wheat scab

2001

2017

On the ring rot of pears and the causal organism, especially on its perfect generation Physalospora piricola n. sp

1933

The Botryosphaeriaceae: genera and species known from culture

2013

Plant disease diagnosis

1997

A new apple tree canker

1919

Compendium of fruit tree pests in China

Botryosphaeria and Physalospora on courrant and apple

1925

Combined multiple gene genealogies and phenotypic characters differentiate several species previously identified Botryosphaeria dothidea

2004

Confronting the constraints of morphological taxonomy in the Botryosphaeriales

2014

Botryosphaeria eucalyptorum sp. nov., a new species in the B. dothidea-complex on Eucalyptus in South Africa

2001

2014

Phylogenetic and pathogenic analyses show that the causal agent of apple ring rot in China is Botryosphaeria dothidea

2012

... 该病于1929年在辽宁发现(景学富等 1963).目前在我国大部分苹果产区普遍发生，尤其在渤海湾苹果产区的河北、山东、辽宁等省和黄河故道区河南和江苏省等地严重发生(胡清玉等 2016).2001年烟台地区苹果轮纹病大发生，一般果园烂果率在30%-50%，部分果园甚至绝收(董娟华和李保华 2009).随着苹果套袋技术的推广，果实发病得到有效控制，但用药的减少导致枝干发病愈趋严重(Tang et al. 2012).近些年，随着气候、栽培制度及大量异地苗木调运等原因，该病在黄土高原区的陕西、山西、甘肃省矮化果园危害也在加重，幼树大面积死亡现象常有发生. ...

... 苹果轮纹病可以侵染果实和枝干，引起的症状也有很大变化，病原复杂多样，病原菌的分类系统也在一直变化，导致该病害的鉴定非常困难.在我国，对该类病害的定义存在异议，导致在不同专著中的描述差异很大.一些著作作为一种病害，称为苹果轮纹病或苹果干腐病(冷怀琼1987;方中达1996;Tang et al. 2012;侯明生和黄俊斌2014;国立耘等2020)，或苹果和梨轮纹病(高学文和陈孝仁2018);一些著作作为2种病害：苹果轮纹病和苹果干腐病(浙江农业大学等1978;中国农作物病虫害编委会1979;中国农业科学院果树研究所1994;刘克均1996;张志铭1996;中国农业科学院植物保护研究所1996;王江柱等2018);还有一些描述为3种病害，分别为苹果轮纹病、苹果干腐病和果实轮纹病(或轮纹烂果病) (江苏农科院植物保护系1997;王江柱和解金斗2019;鲁传涛2021).对病原学名的使用也较混乱，如苹果干腐病的病原有贝林格葡萄座腔菌B. berengeriana、贝克葡萄座腔菌B. bakeri、葡萄座腔菌B. dothidea及茶藨子葡萄座腔菌B. ribis等，苹果轮纹病的病原有贝林格葡萄座腔菌梨生专化型B. berengeriana f. sp. piricola、葡萄座腔菌B. dothidea、轮纹大茎点M. kuwatsukai及梨生囊壳孢Physalospora piricola等(表1). ...

Unraveling Colletotrichum species associated with Glomerella leaf spot of apple

2019

1954

A re-evaluation of the bitunicate Ascomycetes with keys to families and genera

1975

Comparative genomics of Botryosphaeria dothidea and B. kuwatsukai, causal agents of apple ring rot, reveals both species expansion of pathogenicity-related genes and variations in virulence gene content during speciation

2018

... 由于B. dothidea和B. kuwatsukai的ITS、ACT、tubulin等片段的核苷酸差异很小，对2个种的区别力不足，因此对病原菌的多基因系统发育分析与鉴定中，需要选择信息化差异基因，如tef、His等.Wang et al. (2018)测序并比较分析了B. dothidea和B. kuwatsukai全基因组序列，发现后者丢失了一系列次级代谢合成关键酶、碳水化合物活性酶和植物细胞壁降解酶等致病相关基因，为相关差异致病基因的功能研究提供一些指向，建议加强相关差异致病功能基因及致病机制的解析. ...

Comparison of mitochondrial genomes provides insights into intron dynamics and evolution in Botryosphaeria dothidea and B

2021

Illustrated handbook of diagnosis and prevention of pests and diseases on deciduous fruit tree

2018

2019

A secreted lignin peroxidase required for fungal growth and virulence and related to plant immune response

2022

Multiple locus genealogies and phenotypic characters reappraise the causal agents of apple ring rot in China

2015

... . 2015). ...

Effects of environmental factors on internal bark necrosis of apple trees

2008

Species of the genera of Glomerella and Guignardia with special reference to their imperfect stages

1961

Evaluating species in Botryosphaeriales

2021

Toward a natural classification of Botryosphaeriaceae: a study of the type specimens of Botryosphaeria sensu lato

2021

Pome fruits canker

1996

1978

Botryosphaeria rosaceae sp. nov. and B. ramosa, new botryosphaeriaceous taxa from China

2017

果树病害各论

1989

リンコ粗皮病及ぴいば皮病病斑より分离てれゐ系状菌

1980

リンコ果实腐败症の病原菌について

1984

... 均可以引起果实的腐烂症状，在一些苹果上表现为轮纹症状，但是两种病原引起的症状没有差异(小金泽硕城和佐久间勉1984;陈策1993;Xu et al. 2015). ...

苹果干腐病和轮纹病的发生与防治

1993

... ;陈策1993;Xu et al. 2015). ...

苹果果实轮纹病研究进展

1999

中国苹果病害病原菌物名录

2021

苹果轮纹病的研究进展

2009

1996

苹果和梨轮纹病

2018

中国苹果枝干轮纹病发生和防治情况

2009

... 苹果轮纹病(apple ring rot)严重危害果实和枝干，导致苹果果实腐烂、枝干表皮疣突、溃疡，危害幼树导致整树枯死，与苹果树腐烂病和早期落叶病被并称为我国苹果产业的三大病害(国立耘等 2009;李保华等 2013). ...

2020

2014

中国苹果病害发生与分布现状调查

2016

1997

苹果干腐病发生与防治

1963

1987

我国苹果主要病害研究进展与病害防治中的问题

2013

仁果轮纹病

1996

2021

2001

2018

2019

环境因子对苹果粗皮病发生的影响

2008

... 苹果轮纹病引起疣皮症状，大量疣状突起联合、树皮开裂最终表现“粗皮”症状.苹果树锰过剩也可以引起粗皮病，称为苹果粗皮病(internal bark necrosis, IBN)或苹果生理粗皮病.苹果生理粗皮病多数在枝条的光滑表皮上形成疹状突起，少量也在皮孔部位成疹状突起.粗皮病果园枝条韧皮部锰含量均超过200 mg/kg，个别达到600 mg/kg以上(徐圣友等 2008).生理粗皮病在我国多发生于沿海地区，山东胶东半岛发生较为严重，该地区病原性粗皮也普遍发生.在苹果粗皮病防治过程中，需要明确是锰毒害引起的生理性病害还是苹果轮纹病引起的病原性病害.锰过多是否会加重苹果轮纹病的发生也值得进一步深入研究. ...

仁果干腐病

1996

1978

1994

1996

1979

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