热带作物学报

香橼感染黄龙病菌症状表现及病原分布

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廖思雨 , 吴晓雪 , 刘鑫慧 , 谢桃 , 郑丽霞 , 陈文胜 ^* , 李桃 ^*

热带作物学报 | 植物保护与生物安全 2025,46(4): 987-995

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热带作物学报 |植物保护与生物安全 2025 , 46 (4) : 987 -995

香橼感染黄龙病菌症状表现及病原分布

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廖思雨（2000—），女，硕士研究生，研究方向：柑橘黄龙病病原。

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廖思雨（2000—），女，硕士研究生，研究方向：柑橘黄龙病病原。

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廖思雨, 吴晓雪, 刘鑫慧, 谢桃, 郑丽霞, 陈文胜^*, 李桃^*

作者信息

佛山大学农业与生物工程学院，广东佛山　528225

通讯作者:

^* 陈文胜（CHEN Wensheng），E-mail：505827333@qq.com

李桃（LI Tao），E-mail：litao@fosu.edu.cn。

Symptoms and Distribution Patterns of Citrus medica Infected with Candidatus Liberibacter Asiaticus

Siyu LIAO, Xiaoxue WU, Xinhui LIU, Tao XIE, Lixia ZHENG, Wensheng CHEN^*, Tao LI^*

Affiliations

School of Agricultural and Biological Engineering, Foshan University, Foshan, Guangdong 528225, China

出版时间: 2025-04-25 doi: 10.3969/j.issn.1000-2561.2025.04.021

文章导航

摘要

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香橼（Citrus medica）为柑橘属较古老的栽培品种之一，也是现今具有重要商业价值的柑橘类水果的亲本或祖先。香橼因其营养价值和药用价值在柑橘产业中占据重要地位，我国的西南部为香橼起源和多样性的重要中心，香橼种植具有一定规模。柑橘黄龙病（Citrus Huanglongbing）在我国的柑橘主要产区已蔓延数百年，香橼受柑橘黄龙病为害严重，但目前尚无系统描述香橼感染黄龙病后的表现症状及组织解剖结构等的相关研究。本研究以感染黄龙病香橼及健康香橼为材料，观察感染黄龙病香橼叶片及果实等发病症状，并对柑橘黄龙病菌（Candidatus Liberibacter spp.）在香橼不同组织部位的分布规律进行探究，同时对染病香橼及健康香橼组织解剖结构及多项生理指标进行分析与测定。结果表明：染病香橼叶片表现为典型的黄化、斑驳及花叶等症状，一些枝条呈现丛枝、异常开花现象，但未观察到“红鼻子果”。柑橘黄龙病菌在同一香橼植株不同部位的分布不均匀，在橘络中含量最高，老叶叶中脉次之，须根未检测到柑橘黄龙病菌。染病香橼的韧皮部、木质部、薄壁组织及根部细胞排列不规则，韧皮部组织形变坍塌，与健康香橼植株相比，染病香橼主根韧皮部厚度显著下降（P<0.05），在染病香橼各组织切片中均未观察到明显淀粉粒积累现象，但对淀粉含量测定时发现，染病香橼老叶、新叶、主根、须根和橘络中的淀粉含量均高于对应健康组织。染病香橼植株各组织的过氧化氢含量显著增加（P<0.05），尤其是在幼嫩组织（新叶、须根）中显著积累；与健康香橼植株相比，染病香橼植株组织，尤其是主根与老叶叶中脉胼胝质沉积现象更明显。本研究结果可为今后探究柑橘黄龙病菌对柑橘侵染造成的破坏特点，以及柑橘对柑橘黄龙病菌的防御机制等相关研究提供理论依据。

关键词

香橼 / 柑橘黄龙病 / 症状 / 病原分布

Abstract

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Citrus medica is an older cultivated varieties of the genus Citrus and the parent or ancestor of Citrus fruits with significant commercial value. Due to its nutritional and medicinal values, C. medica occupies an important position in the citrus industry. The southwestern part of China is an important center for the origin and diversity of C. medica, and its cultivation has reached a certain scale. Citrus Huanglongbing (HLB) has been spreading for hundreds of years in the major citrus-producing areas in China. C. medica infected with HLB and healthy C. medica were used as the materials. Firstly, the symptoms of the leaves and fruits of C. medica infected with HLB were observed. Secondly, the distribution pattern of HLB in different tissue parts of C. medica was explored. Meanwhile, the tissue anatomical structures and multiple physiological indicators of C. medica infected with HLB and healthy C. medica were analyzed and measured through comparison. Through comparative observation, it was found that the leaves of C. medica infected with HLB showed typical symptoms such as yellowing, mottling and mosaic, and some branches presented symptoms of witches broom and abnormal flowering, but no “red nose fruit” was observed. The distribution of HLB in different parts of the same C. medica was uneven, with the highest content in the fruit pith, followed by the midrib of the old leaves, and HLB was not detected in the fibrous roots. In C. medica infected with HLB, the cells in the phloem, xylem, parenchyma and root were arranged irregularly, the phloem tissue was deformed and collapsed, and the thickness of the phloem of the taproot was significantly decreased compared with that of the healthy C. medica at the same period (P<0.05). No obvious starch grain accumulation was observed in each tissue section of C. medica infected with HLB, but when the starch content was measured, it was found that the starch contents in the old leaves, new leaves, taproot, fibrous roots and fruit pith of C. medica infected with HLB were all higher than those of the corresponding healthy tissues. The hydrogen peroxide content in each tissue of the diseased C. medica increased significantly (P<0.5), especially significantly accumulated in the young tissues (new leaves, fibrous roots) infected with HLB. Compared with healthy C. medica, the callose deposition phenomenon in the tissues of C. medica infected with HLB, especially in the main root and the midrib of the old leaves, was more obvious. The research results of this study could provide a theoretical basis for the relevant studies on exploring the characteristics of damage caused by HLB and the defense mechanism of citrus against HLB in the future.

Key words

Citrus medica / Citrus Huanglongbing / symptoms / bacterial distribution

引用本文

廖思雨, 吴晓雪, 刘鑫慧, 谢桃, 郑丽霞, 陈文胜, 李桃. 香橼感染黄龙病菌症状表现及病原分布. 热带作物学报, 2025 , 46 (4) : 987 -995 . DOI: 10.3969/j.issn.1000-2561.2025.04.021

Siyu LIAO, Xiaoxue WU, Xinhui LIU, Tao XIE, Lixia ZHENG, Wensheng CHEN, Tao LI. Symptoms and Distribution Patterns of Citrus medica Infected with Candidatus Liberibacter Asiaticus[J]. Chinese Journal of Tropical Crops, 2025 , 46 (4) : 987 -995 . DOI: 10.3969/j.issn.1000-2561.2025.04.021

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柑橘黄龙病（Citrus Huanglongbing，HLB）是由α-变形菌纲韧皮部杆菌暂定属细菌（Candidatus Liberibacter spp.）引起且对柑橘生产具有毁灭性的病害，对世界柑橘产业造成巨大的经济损失^[1]。目前基于16S rRNA基因位点的细菌分类体系及病原菌的热敏感性、流行区域及昆虫媒介等特点，将柑橘黄龙病菌分为柑橘黄龙病菌亚洲种（Candidatus Liberibacter asiaticus，CLas）、柑橘黄龙病菌美洲种（Candidatus Liberibacter americanus，CLam）及柑橘黄龙病菌非洲种（Candidatus Liberibacter africanus，CLaf），在我国仅发现了柑橘黄龙病菌亚洲种^[2]。柑橘黄龙病在我国为害已长达百年之久，早在1919年REINKING在我国南方地区进行病害调查时发现了一些柑橘叶片上出现了典型的斑驳黄化症状，这一记载也被广大学者普遍认为是柑橘黄龙病在我国出现的最早证据^[3]。在感染柑橘黄龙病菌后，柑橘叶片会停止转绿，叶片逐渐变黄、革质化，叶片无光泽，一般会出现黄化、花叶、斑驳及绿岛等症状，一些植株也会出现丛枝现象，果实发育畸形，种子败育，一些柑橘品种果实染病后会表现典型的“红鼻子果”；在发病后期，柑橘植株根系完全腐烂，大量叶片脱落，直至植株枯死^[3-5]。

柑橘黄龙病为系统性侵染病害，但其病原菌在寄主组织内分布不均匀。对一些感染柑橘黄龙病菌的柑橘品种不同组织部位的病原浓度进行定量发现，柑橘黄龙病菌在不同部位的浓度差异很大。如染病沙田柚花器和种子，柑橘黄龙病菌在种子种皮中的浓度远远高于花器的雌蕊、雄蕊及花粉等组织^[6]。在染病贡柑中，果实橘络中的柑橘黄龙病菌浓度则显著高于叶片、果轴等部位；在染病砂糖橘中，橘络中的柑橘黄龙病菌浓度最高，其余依次为果皮、中心柱、近果处叶片等部位，并且柑橘黄龙病菌在染病柑橘橘络中分布不均匀，倾向定殖于橘络的中间区域^[7-10]。柑橘感染柑橘黄龙病菌后，不同症状的组织所含的柑橘黄龙病菌浓度也有差异，张培等^[11]对感染柑橘黄龙病菌的砂糖橘“红鼻子果”、斑驳叶片、黄化叶片及无症状果实中的柑橘黄龙病菌浓度进行定量分析发现，柑橘黄龙病菌在“红鼻子果”中含量最高；陈燕玲等^[12]的研究结论与之类似；另外黄桢辉等^[13]对砂糖橘、W-默科特、红心蜜柚和沙田柚的不同生长时期果枝中柑橘黄龙病菌进行定量分析发现，柑橘黄龙病菌在果实的橘络、果中轴及种皮处富集。除病原菌在寄主组织分布不均匀外，一些柑橘品种在染病后表现的症状也存在一些差异。如广佛手感染柑橘黄龙病菌后叶片虽然会出现典型的斑驳黄化症状，部分病果畸形且小，但未出现“红鼻子果”症状，并且柑橘黄龙病菌并未在成熟果实中富集^[14-15]。

香橼（Citrus medica）是柑橘主要栽培品种之一，也是商业上重要柑橘品种的亲本或祖先^[16]。香橼主要产区为中国西南部以及西藏东南部，在印度、地中海地区和加勒比海沿岸国家也有广泛种植^[17]。柑橘在中药、观赏用途和香料制剂中有着广泛的应用，香橼果实中存在丰富的有益于人类健康的生物活性化合物，使其成为开发健康食品及药材的理想原料^[17-19]。柑橘黄龙病会致使香橼植株出现花期异常、果实畸形脱落、植株枯死等症状，严重影响香橼的果实品质及经济效益。现阶段有关香橼感染柑橘黄龙病菌后的生理指标变化及病原分布规律尚无系统报道，因此本研究对感染柑橘黄龙病菌的香橼不同组织部位的症状表现、病原分布、组织解剖结构及相关生理指标进行系统测定，解析黄龙病菌致病表现及对寄主组织的破坏特点，为后续黄龙病菌的致病机制研究提供理论基础。

1　材料与方法

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1.1　材料

于2024年10月在广东省佛山市佛山大学农业与生物工程学院实验基地开展试验，选取2年生健康香橼植株栽种于温室大棚单独培养，选取部分幼苗嫁接携带柑橘黄龙病菌的接穗，2 a后待嫁接携带柑橘黄龙病菌接穗的香橼黄龙病菌检测为阳性、植株表现症状且枝条挂果后，对健康香橼与感染柑橘黄龙病菌且具有典型症状的香橼不同组织进行采集。如图1所示，将组织分为新叶叶肉、新叶叶脉、新叶叶柄、老叶叶肉、老叶叶脉、老叶叶柄、果蒂、橘络、主干韧皮部、主根、须根共11个部分。样品采集后经液氮速冻后，置于–80 ℃超低温冰箱保存备用。用于胼胝质测定的新鲜组织材料采集后低温保存即刻带回实验室进行测定。

1.2　方法

1.2.1　香橼组织DNA提取与柑橘黄龙病菌的定量分析

分别称取香橼各组织100 mg，充分研磨后采用OMEGA BIO-TEK公司的HP Plant DNA Kit（2485-02）植物DNA提取试剂盒对香橼各组织总DNA进行提取。利用超微量紫外-可见光分光光度计NanoDrop one（Thermo Fisher Scientific）对提取的DNA纯度与浓度进行测定，然后置于–20 ℃保存备用。采用实时荧光定量PCR技术对上述11个组织部位DNA中的柑橘黄龙病菌数量进行定量分析，具体检测方法参考BAO等^[20]的方法，对所有组织DNA样品进行SYBR Green实时荧光定量PCR检测，采用两步法程序，95 ℃变性5 min；95 ℃退火10 s，60 ℃延伸30 s，40个循环。以香橼各组织的DNA为模板，并以灭菌ddH₂O为空白对照，以健康香橼组织DNA为阴性对照，以已知含柑橘黄龙病菌的香橼组织所提取DNA为阳性对照，进行荧光定量PCR检测。待反应结束后，根据得到的CT值判断样品是否含有柑橘黄龙病菌（CT<32时，则判定该样品携带柑橘黄龙病菌）。利用含有引物目的片段不同浓度梯度的重组质粒制作标准曲线，得到标准方程后，对香橼各组织DNA中的柑橘黄龙病菌浓度进行量化。柑橘黄龙病菌浓度以每纳克（ng）DNA含有的柑橘黄龙病菌拷贝数表示。

1.2.2　香橼组织解剖结构观察与厚度测量

采集的健康与感染柑橘黄龙病菌的香橼植株组织利用固定液固定24 h后，将组织取出修平。将香橼组织依次进行脱水浸蜡、样品包埋、切片、番红染色、固绿染色等处理，待处理结束后利用中性树胶将组织切片封片，得到的香橼组织切片利用显微镜进行观察，并对图像进行采集分析。

利用尼康光学显微镜（Nikon ECLIPSE Ni-U）及NIS element viewer 4.20软件对上述处理后的香橼组织石蜡切片进行观察，设定标尺后对视野清晰度进行矫正后拍照保存，每个样品切片各5个重复。利用ImageJ软件对切片照片中的各组织厚度进行测量，每个组织观察5个玻片，并对每个玻片的目的组织进行20次测量，组织厚度取平均值。

1.2.3　香橼组织生理指标测定

（1）不同组织部位淀粉含量。分别称取健康香橼和感病香橼植株不同部位样品各0.03 g，在液氮条件下将其磨碎，然后根据北京索莱宝科技有限公司的淀粉含量检测试剂盒说明书进行提取与测定。

（2）不同组织部位过氧化氢含量。将采集的植物样品用磷酸盐缓冲液（PBS）擦洗干净，再用吸水纸擦干，然后剪碎放入研钵，液氮条件下磨成粉，称取植物粉末0.1 g加入1.5 mL PBS，涡旋振荡1 min，按12 000 r/min离心10 min取上清液。过氧化氢指标检测所使用的试剂盒为96样微板法，购自南京建成生物工程研究所，所用设备为赛默飞的全自动酶标仪。

（3）不同组织部位胼胝质沉积情况观察。分别采集健康香橼和感病香橼植株的新叶叶脉、老叶叶脉、老根和主干韧皮部。将新鲜香橼组织切成2 mm左右的薄片，直接浸于AFF固定液中24 h。使用无水乙醇浸洗2次，每次1 min，然后转入至少10倍体积的100%乙醇中保存；染色前取出组织，浸于50%乙醇中平衡30 min，取出稍沥干。随后置于1×PBS缓冲液中平衡30 min，取出稍沥干。使用苯胺蓝对香橼不同组织进行避光染色1 h，利用奥林巴斯显微镜（OLYMPUS BX53）对上述处理后的香橼组织进行观察。以上生理指标测定均5个生物学重复。

1.3　数据处理

利用SPSS Statistics 25.0软件（IBM Corp.，Armonk，N.Y.，USA）对试验数据进行统计与分析，利用Origin 2018软件（OriginLab Corp.，Northampton，MA 01060，USA）对数据结果进行可视化。

2　结果与分析

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2.1　染病香橼植株症状表现

在嫁接携带柑橘黄龙病菌的接穗2 a后，选取柑橘黄龙病菌检测呈阳性且表现症状的香橼植株进行采样观察。结果显示，与健康香橼叶片相比，染病香橼叶片出现叶肉组织部分或均匀黄化、斑驳、叶中脉黄化、花叶及绿岛等典型柑橘黄龙病症状；染病叶片呈现狭小、细长及革质化状态，未观察到缺锌症状（图2A，图2B）；一些叶片严重黄化的枝条后续叶片会大量脱落，直至枝条干枯死亡，同时也观察到丛枝现象（图2D~图2F）。染病的果实出现果蒂处黄化、果体变小、畸形及不均匀黄化现象，尚未观察到“红鼻子果”症状（图2C），在冬季观察到一些染病植株出现异常开花现象（图2G）。

2.2　黄龙病菌在香橼不同组织的分布

通过对染病香橼植株的11个部位进行柑橘黄龙病菌检测发现，各组织均检测到柑橘黄龙病菌（图3）。实时荧光定量PCR结果表明，柑橘黄龙病菌在染病香橼植株中分布不均匀，其中橘络的柑橘黄龙病菌浓度最高，老叶叶脉次之。柑橘黄龙病菌在染病香橼植株不同组织的浓度从高到低依次为：橘络（1622.13 cells/ng）>老叶叶脉（521.97 cells/ng）>果蒂（380.55 cells/ng）>老叶叶柄（319.93 cells/ng）>主干韧皮部（236.46 cells/ng）>主根（105.64 cells/ng）>新叶叶柄（43.10 cells/ng）>新叶叶脉（29.45 cells/ng）>老叶叶肉（27.49 cells/ng）>新叶叶肉（1.42 cells/ng）>须根（0.14 cells/ng）。

2.3　感染黄龙病后香橼不同组织的解剖结构变化

通过光学显微镜对健康香橼和染病香橼植株的新老叶片、主根及须根的组织石蜡切片进行观察发现，染病香橼和健康香橼的各组织结构均比较清晰，细胞均清晰可见。其中健康香橼的新叶与老叶叶中脉薄壁组织、韧皮部及木质部细胞排列整齐规则（图4A，图4C），而染病香橼新叶与老叶叶中脉薄壁细胞和韧皮部细胞的整齐度下降，细胞排列不规则，韧皮部发生严重形变，呈现组织塌陷现象，在薄壁组织、韧皮部和髓部淀粉粒积累的现象不明显（图4B，图4D）；通过对比发现，健康香橼新叶与老叶叶肉的栅栏组织排列紧密整齐，海绵组织空隙小（图4E，图4G）；染病香橼新叶与老叶叶肉的栅栏组织细胞排列整齐度降低，海绵组织空隙大，部分海绵组织细胞被破坏（图4F，图4H）。染病香橼主根的韧皮部与木质部细胞排列不规则，大量薄壁组织、韧皮部与木质部细胞形态变化，在染病的须根中也观察到类似现象（图4I~图4L）。

对健康香橼与染病香橼叶片、主根和须根的各组织厚度进行测量比较发现，与健康香橼植株相比，染病香橼植株的新老叶片和须根的韧皮部组织厚度有所下降，但差异不显著；染病香橼植株的叶片、主根和须根的木质部、薄壁组织厚度均显著下降（P<0.05，图5A~图5D）。染病香橼的新叶和须根各组织厚度的下降幅度均低于染病老叶和主根各组织厚度，如在染病香橼中，主根的韧皮部组织厚度为110.26 μm，健康香橼主根韧皮部组织厚度为350.49μm，二者差异显著（P<0.05），健康香橼主根韧皮部厚度为染病香橼的3.18倍（图5C），而健康香橼与染病香橼须根中的韧皮部组织厚度无显著差异（图5D），这一结果表明柑橘黄龙病菌对香橼幼嫩组织的破坏程度低于成熟组织。

2.4　感染黄龙病后香橼不同组织的生理指标变化

对感染柑橘黄龙病菌后香橼不同组织的淀粉含量进行测定，结果发现，染病香橼的新叶、老叶、主根及橘络中的淀粉含量均显著高于健康香橼（P<0.05），其中染病香橼老叶叶肉的淀粉含量较健康香橼上升93%，其上升幅度高于染病香橼新叶叶肉中淀粉的积累速度，并且染病香橼老叶叶肉中的淀粉含量最高，橘络中积累最少（图6）。

通过对感染柑橘黄龙病菌后香橼不同组织过氧化氢含量进行测定，结果显示，除老叶叶脉外，染病植株各组织的过氧化氢含量均显著高于健康香橼（P<0.05），并且感染柑橘黄龙病菌后新叶（叶肉、叶脉）的过氧化氢含量也高于老叶（叶肉、叶脉）及主根（图7）。

通过对健康香橼与染病香橼的新叶叶脉、老叶叶脉、主干韧皮部和主根进行苯胺蓝染色，在荧光显微镜的激发光下，观察各组织的韧皮部胼胝质沉积情况。结果发现，健康香橼的各组织韧皮部细胞中胼胝质沉积较少；而染病香橼老叶叶脉、主干韧皮部和老根处韧皮部均可观察到明显的荧光染色，说明染病香橼的老叶叶脉、主干及老根的韧皮部出现了明显的胼胝质沉积现象（红色箭头指向为胼胝质），而在染病香橼新叶中，沉积现象不明显（图8）。

3　讨论

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本研究在对感染柑橘黄龙病菌的香橼叶片及果实症状进行观察时，未发现缺锌症状，并且未观察到典型的“红鼻子果”症状，这与罗晓玲等^[15]在感染柑橘黄龙病菌广佛手叶片上的表现症状类似，而像砂糖橘、贡柑及茶枝柑等品种在感染柑橘黄龙病菌后其叶片极易观察到缺锌及“红鼻子果”症状^[7,10,12]，推测这一现象可能与不同柑橘品种有关。另外通过测定柑橘黄龙病菌在香橼不同组织中的含量时发现，柑橘黄龙病菌在香橼橘络中含量最高，与前人研究^[7-10,12]相似，因此后期可以香橼橘络为材料获取高浓度柑橘黄龙病菌。柑橘茎尖组织常作为柑橘脱毒技术的植物材料^[21]，在本研究中染病香橼的须根柑橘黄龙病菌检测呈阴性，因此后续若对一些稀有香橼种质资源进行保存、扩繁，可尝试利用香橼的茎尖组织进行繁殖。

在本研究中，对感染柑橘黄龙病菌香橼不同组织厚度进行测量时发现，香橼的新叶、老叶及须根的韧皮部组织厚度未受柑橘黄龙病菌侵染的影响，染病香橼与健康香橼的韧皮部组织厚度差异不显著，在对不同组织的切片进行观察也未发现大量的淀粉粒沉积现象，前人研究表明佛手品种的柑橘对柑橘黄龙病具有一定的抗性^[22]，因此推测染病香橼这几个组织的韧皮部厚度与健康香橼相比差异不大，可能也与香橼对柑橘黄龙病具有一定的抗性有关。在对淀粉含量进行测定时发现，在染病香橼中老叶叶肉淀粉含量最多，橘络最少，前人研究表明，柑橘黄龙病菌侵染后会导致叶片碳水化合物运输异常^[23]，同时老叶积累光合产物较多，橘络一般运输水分和养分，并非淀粉储存的主要部位，因此淀粉含量在染病老叶中高于其他组织，而橘络中较低。

柑橘感染柑橘黄龙病菌后，会激活自身免疫反应，其中包括诱导一些活性氧，如过氧化氢及胼胝质合成等与防御相关基因的表达，以应对柑橘黄龙病菌的侵染^[24]。在本研究中，香橼幼嫩组织（新叶）感染柑橘黄龙病菌后，过氧化氢含量显著上升，推测这一现象的出现可能是由于香橼幼嫩组织防御组织不发达，如缺乏厚的细胞壁及角质层，且代谢活跃，由此香橼幼嫩组织对柑橘黄龙病菌侵染比较敏感，活性氧含量显著上升；胼胝质与柑橘防御反应相关^[23-24]，柑橘感染柑橘黄龙病菌后会出现胼胝质沉积的现象，本研究发现染病香橼的各组织胼胝质沉积较多，而在健康香橼组织较少，与前人研究结果^[25]一致。综上所述，本研究对感染柑橘黄龙病菌后香橼植株的外观症状表现、组织细胞形态、病菌分布及生理指标变化进行了深入探究解析，研究结果可为后续黄龙病菌致病机制的研究及柑橘黄龙病防治提供理论依据。

基金

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国家自然科学基金青年基金项目(32202277)

参考文献

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文献

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2025年第46卷第4期

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doi: 10.3969/j.issn.1000-2561.2025.04.021

接收时间：2024-12-07
首发时间：2026-06-24
出版时间：2025-04-25

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收稿日期：2024-12-07
录用日期：2024-12-30

基金

国家自然科学基金青年基金项目(32202277)

作者信息

佛山大学农业与生物工程学院，广东佛山　528225

通讯作者:

^* 陈文胜（CHEN Wensheng），E-mail：505827333@qq.com

李桃（LI Tao），E-mail：litao@fosu.edu.cn。

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2种不同金属材料的力学参数

科 Family	属数 Number of genus	种数 Number of species	占总种数比例 Percentage of total species (%)	属 Genus	种数 Number of species	占总种数比例 Percentage of total species (%)
鹅膏菌科Amanitaceae	2	11	5.26	鹅膏菌属 Amanita	10	4.78
小菇科 Mycenaceae	2	12	5.74	丝盖伞属 Inocybe	5	2.39
多孔菌科 Polyporaceae	8	14	6.70	蜡蘑属 Laccaria	5	2.39
红菇科 Russulaceae	3	23	11.00	小皮伞属 Marasmius	6	2.87
				小菇属 Mycena	11	5.26
				光柄菇属 Pluteus	5	2.39
				红菇属 Russula	17	8.13
				栓菌属 Trametes	5	2.39

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