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Article(id=1195814097355849845, tenantId=1146029695717560320, journalId=1190317699101192196, issueId=1195814096831561845, articleNumber=1001-2494(2024)23-2199-06, orderNo=null, doi=10.11669/cpj.2024.23.001, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1715875200000, receivedDateStr=2024-05-17, revisedDate=null, revisedDateStr=null, acceptedDate=null, acceptedDateStr=null, onlineDate=1763033897960, onlineDateStr=2025-11-13, pubDate=1733587200000, pubDateStr=2024-12-08, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1763033897960, onlineIssueDateStr=2025-11-13, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1763033897960, creator=13701087609, updateTime=1763033897960, updator=13701087609, issue=Issue{id=1195814096831561845, tenantId=1146029695717560320, journalId=1190317699101192196, year='2024', volume='59', issue='23', pageStart='2199', pageEnd='2298', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=0, createTime=1763033897836, creator=13701087609, updateTime=1763034755097, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1195817692507718044, tenantId=1146029695717560320, journalId=1190317699101192196, issueId=1195814096831561845, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1195817692507718045, tenantId=1146029695717560320, journalId=1190317699101192196, issueId=1195814096831561845, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=2199, endPage=2204, ext={EN=ArticleExt(id=1195814097590730871, articleId=1195814097355849845, tenantId=1146029695717560320, journalId=1190317699101192196, language=EN, title=Comparative Research Progress of Zhushiling and Jishiling, columnId=null, journalTitle=Chinese Pharmaceutical Journal, columnName=null, runingTitle=null, highlight=null, articleAbstract=

Polyporus umbellatus, a traditional medicinal fungus in China, is used medicinally for its sclerotium. It is commonly categorized into two types, Zhushiling and Jishiling, based on the morphology of the sclerotia. Cultivation of Polyporus umbellatus heavily relies on the asexual reproduction of its sclerotia and the absence of clear distinction between Zhushiling and Jishiling often results in the confusion and intermixing of seed sclerotia. This can lead to the instability in quality and yield of Polyporus umbellatus medicinal materials, thereby adversely affecting the robust development of the Polyporus umbellatus industry. This article provides a comparative overview of Zhushiling and Jishiling in 4 aspects: morphology, quality, genetics and spawn cultivation. The results indicate that although the differentiation between Zhushiling and Jishiling based on morphological features such as sclerotium branches and surface characteristics holds certain significance, there remains a necessity to develop more scientific and precise criteria for this classification. The traditional perspective that Zhushiling is of superior quality compared with Jishiling is often based on empirical observations rather than rigorous scientific evidence. It is imperative to conduct comprehensive and systematic studies to objectively evaluate the quality and medicinal properties of both types. The variation in the sclerotial morphology of Polyporus umbellatus may be the result of a complex interplay among genetic, geographic, environmental factors, as well as other factors that have yet to be identified. Key technological breakthroughs in cultivation of the Polyporus umbellatus spawn leading to the formation of the sclerotia have not yet been achieved. However, the strains of Jishiling may possess greater potential for the development of spawn cultivation techniques. Zhushiling and Jishiling exhibit differences across 4 distinct aspects. Conducting the systematic analysis of Polyporus umbellatus from various perspectives is significantly important for the evaluation of its resources and for maximizing its potential value.

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猪苓为我国传统药用真菌,以菌核入药,根据菌核形态常将其分为猪屎苓和鸡屎苓。猪苓栽培依赖种苓无性繁殖,人们对猪屎苓和鸡屎苓的认知不足往往造成苓种混杂,从而导致猪苓药材质量和产量不稳定,影响猪苓产业的健康发展。本文从形态、品质、遗传和菌种栽培4个方面对猪屎苓和鸡屎苓进行了比较与综述。结果表明,虽然以菌核分支、表面特性等形态特征区分猪屎苓与鸡屎苓具有一定的意义,但仍需要建立更科学的分类标准;传统认为猪屎苓质量优于鸡屎苓的观点缺乏科学性,需进行更系统的评价;猪苓菌核形态的差异可能是由遗传、地理、环境或多种因素共同作用的结果;猪苓菌种栽培形成菌核关键技术还未完全突破,鸡屎苓菌株可能更具有开展菌种栽培的潜力。猪屎苓和鸡屎苓在这4个方面均存在差异,从不同角度对猪苓进行系统分析,对于评估猪苓资源并实现其价值的最大化具有重要意义。

, correspAuthors=郭顺星, authorNote=null, correspAuthorsNote=
* 郭顺星,男,博士,教授 研究方向:药用植物内生真菌生物学 Tel:(010)57833259
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刘幽言,女,硕士研究生 研究方向:药用菌物学

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刘幽言,女,硕士研究生 研究方向:药用菌物学

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刘幽言,女,硕士研究生 研究方向:药用菌物学

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指标 猪屎苓 鸡屎苓
形态[9,17-20] 猪屎状、瘤状、长形块状 鸡爪状、鸡屎状、条状、片状、姜状、饼状
个头[11] 大 小
分枝[14-16] 分枝少、粗壮而短 分枝多且细
表面[15-16] 光滑,有瘤状突起 褶皱明显、凹陷深
产量[11-12] 高 低
土壤条件[20] 肥沃 贫瘠
同一窝中分布[20] 多分布在下层 多分布在上层
收购价格[7] 高 低
市场流通量[13] 高 低
地域分布[19] 华北、西北地区居多 东北长白山区居多
菌种栽培潜能 鸡屎苓菌株可能更适宜用作菌种栽培猪苓
), ArticleFig(id=1196079280070378118, tenantId=1146029695717560320, journalId=1190317699101192196, articleId=1195814097355849845, language=CN, label=表1, caption=

猪屎苓和鸡屎苓的差异

, figureFileSmall=null, figureFileBig=null, tableContent=
指标 猪屎苓 鸡屎苓
形态[9,17-20] 猪屎状、瘤状、长形块状 鸡爪状、鸡屎状、条状、片状、姜状、饼状
个头[11] 大 小
分枝[14-16] 分枝少、粗壮而短 分枝多且细
表面[15-16] 光滑,有瘤状突起 褶皱明显、凹陷深
产量[11-12] 高 低
土壤条件[20] 肥沃 贫瘠
同一窝中分布[20] 多分布在下层 多分布在上层
收购价格[7] 高 低
市场流通量[13] 高 低
地域分布[19] 华北、西北地区居多 东北长白山区居多
菌种栽培潜能 鸡屎苓菌株可能更适宜用作菌种栽培猪苓
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猪屎苓与鸡屎苓的比较研究进展
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刘幽言 , 李寿建 , 李兵 , 郭顺星 *
中国药学杂志 | 综述 2024,59(23): 2199-2204
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中国药学杂志 | 综述 2024, 59(23): 2199-2204
猪屎苓与鸡屎苓的比较研究进展
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刘幽言, 李寿建, 李兵, 郭顺星*
作者信息
  • 中国医学科学院北京协和医学院药用植物研究所, 北京 100193

    • 刘幽言,女,硕士研究生 研究方向:药用菌物学

通讯作者:

* 郭顺星,男,博士,教授 研究方向:药用植物内生真菌生物学 Tel:(010)57833259
Comparative Research Progress of Zhushiling and Jishiling
Youyan LIU, Shoujian LI, Bing LI, Shunxing GUO*
Affiliations
  • Institute of Medicinal Plant Development, Chinese Academy of Medical Sciences & Peking Union Medical College, Beijing 100193, China
出版时间: 2024-12-08 doi: 10.11669/cpj.2024.23.001
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摘要
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猪苓为我国传统药用真菌,以菌核入药,根据菌核形态常将其分为猪屎苓和鸡屎苓。猪苓栽培依赖种苓无性繁殖,人们对猪屎苓和鸡屎苓的认知不足往往造成苓种混杂,从而导致猪苓药材质量和产量不稳定,影响猪苓产业的健康发展。本文从形态、品质、遗传和菌种栽培4个方面对猪屎苓和鸡屎苓进行了比较与综述。结果表明,虽然以菌核分支、表面特性等形态特征区分猪屎苓与鸡屎苓具有一定的意义,但仍需要建立更科学的分类标准;传统认为猪屎苓质量优于鸡屎苓的观点缺乏科学性,需进行更系统的评价;猪苓菌核形态的差异可能是由遗传、地理、环境或多种因素共同作用的结果;猪苓菌种栽培形成菌核关键技术还未完全突破,鸡屎苓菌株可能更具有开展菌种栽培的潜力。猪屎苓和鸡屎苓在这4个方面均存在差异,从不同角度对猪苓进行系统分析,对于评估猪苓资源并实现其价值的最大化具有重要意义。

猪苓  /  菌核  /  品质  /  遗传多样性  /  菌种栽培

Polyporus umbellatus, a traditional medicinal fungus in China, is used medicinally for its sclerotium. It is commonly categorized into two types, Zhushiling and Jishiling, based on the morphology of the sclerotia. Cultivation of Polyporus umbellatus heavily relies on the asexual reproduction of its sclerotia and the absence of clear distinction between Zhushiling and Jishiling often results in the confusion and intermixing of seed sclerotia. This can lead to the instability in quality and yield of Polyporus umbellatus medicinal materials, thereby adversely affecting the robust development of the Polyporus umbellatus industry. This article provides a comparative overview of Zhushiling and Jishiling in 4 aspects: morphology, quality, genetics and spawn cultivation. The results indicate that although the differentiation between Zhushiling and Jishiling based on morphological features such as sclerotium branches and surface characteristics holds certain significance, there remains a necessity to develop more scientific and precise criteria for this classification. The traditional perspective that Zhushiling is of superior quality compared with Jishiling is often based on empirical observations rather than rigorous scientific evidence. It is imperative to conduct comprehensive and systematic studies to objectively evaluate the quality and medicinal properties of both types. The variation in the sclerotial morphology of Polyporus umbellatus may be the result of a complex interplay among genetic, geographic, environmental factors, as well as other factors that have yet to be identified. Key technological breakthroughs in cultivation of the Polyporus umbellatus spawn leading to the formation of the sclerotia have not yet been achieved. However, the strains of Jishiling may possess greater potential for the development of spawn cultivation techniques. Zhushiling and Jishiling exhibit differences across 4 distinct aspects. Conducting the systematic analysis of Polyporus umbellatus from various perspectives is significantly important for the evaluation of its resources and for maximizing its potential value.

Polyporus umbellatus  /  sclerotia  /  quality  /  genetic diversity  /  spawn cultivation
刘幽言, 李寿建, 李兵, 郭顺星. 猪屎苓与鸡屎苓的比较研究进展. 中国药学杂志, 2024 , 59 (23) : 2199 -2204 . DOI: 10.11669/cpj.2024.23.001
Youyan LIU, Shoujian LI, Bing LI, Shunxing GUO. Comparative Research Progress of Zhushiling and Jishiling[J]. Chinese Pharmaceutical Journal, 2024 , 59 (23) : 2199 -2204 . DOI: 10.11669/cpj.2024.23.001
正文
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猪苓是多孔菌科真菌猪苓[Polyporus umbellatus (Pers.) Fries]的干燥菌核[1],具有利水渗湿等功效。猪苓之名源于其形似猪屎的外观,首次记载于《神农本草经》[2]:“猪苓,一名猳猪矢,味甘平,生山谷。”明代李时珍[3]的《本草纲目》中解释了“苓”字的用意,“猪屎曰零,即苓字,其块零落而下故也”,取“其块零落而下”之意,称猪屎为“零”,而“零”与“苓”通用,所以称为“猪苓”。猪苓的别名如“野猪粪”“枫木苓”“朱苓”等多与其形态特点、生长习性、方音讹传有关[4],自明清开始众多医家多沿用猪苓为其正名。而古籍记载猪苓由“木之余气所结”[3]推测是指猪苓与蜜环菌间特殊的共生关系。经典分类学将猪苓按菌核形态划分[5],普遍认为体形较大,表面光滑、形如猪屎的称为猪屎苓;体形较小,表面多褶皱、形如鸡屎的为鸡屎苓,但目前还缺乏统一的判断标准。
猪苓是我国重要的药用真菌之一,其菌核中含有多糖、甾体、脂肪酸和多酚等多种化学成分,多糖和甾体为主要的药效成分。猪苓多糖具有抗癌、抗病毒、增强免疫力和抗氧化等多种功效,而甾体类化合物在利尿和肾脏保护方面发挥着不可替代的作用[6]。由于猪屎苓和鸡屎苓的判断以菌核表观形态为主要依据,药农仅凭经验选择种苓进行人工栽培往往造成苓种混杂。虽然猪苓菌核的形态遗传比较稳定[7],但生长速度、产量和质量有所差异,苓种混杂会导致猪苓药材质量和产量不稳定,制约猪苓产业的健康可持续发展。故笔者拟从形态、品质、遗传和菌种栽培4个方面对猪屎苓和鸡屎苓进行比较和综述,以促进对猪苓种质的全面了解,为保护和充分利用猪苓种质资源,指导猪苓人工栽培和菌种选育提供理论依据。
1 鸡屎苓和猪屎苓的差异指标
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在实际生产中,药农常根据菌核形态特征将猪苓划分为猪屎苓、鸡屎苓(也称鸡爪苓)[7]、康巴苓[8]、铁蛋苓(也称铁蛋猪屎苓)[8-9]和马屎苓[10]等,其中以猪屎苓和鸡屎苓最为常见和具有代表性。人工栽培猪苓早期,学者以产量区分猪屎苓和鸡屎苓。母猪窝苓,产量大,每窝可挖几十斤;鸡窝苓,产量小,每窝只能挖几斤[11],推测母猪窝苓就是常说的猪屎苓,鸡窝苓就是鸡屎苓。研究者在产量的基础上增加了对猪屎苓和鸡屎苓的体型大小和均匀程度的考究,认为猪屎苓块大、分布不均、大小不等、产量高,多的猪屎苓窝可产40 kg以上;而鸡屎苓的块小、分布均匀、大小基本相等、产量低,最多的鸡屎苓窝不超过5 kg[12]。根据Yang等[13]的市场调查,鸡屎苓的市场流通量远低于猪屎苓,154个商家的猪苓药材中猪屎苓占到96.75%,而鸡屎苓仅占3.25%且均为野生猪苓。
此后学者们主要从分枝、表面光滑程度以及形状3个方面对猪屎苓和鸡屎苓进行描述。猪屎苓分枝小,粗壮而短[14-15],表面较光滑[16],有瘤状突起[15];鸡屎苓分枝多且较细[14,16],表面褶皱明显,凹陷深[15]。在形状方面,研究者普遍认为长形块状[9]、瘤状[17]、形如猪屎的为猪屎苓;条状[18]、片状[19]、形如鸡屎的为鸡屎苓,典型的猪屎苓和鸡屎苓的菌核形态见图1。也有学者提到猪屎苓和鸡屎苓的产地和生长环境不同,华北、西北地区生长多为猪屎苓,东北长白山区生长多为鸡屎苓[19];猪屎苓生长在肥沃的土壤中,而鸡屎苓生长的土壤则更瘠薄。同一窝中两者可能同时存在,鸡屎苓多分布在上层,猪屎苓多分布在下层[20]。本研究根据文献描述,将不同猪苓的差异进行了总结(表1)。
2 猪屎苓和鸡屎苓的质量差异
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在传统中药材评估中,猪屎苓与鸡屎苓的质量评价标准曾主要依赖于外观特征,皮黑肉白,块大体实,无沙石杂质者为上品;皮不太黑,块小,烂碎,肉褐色,褶皱不紧者次之[21]。后人补充了猪苓佳品需新鲜健壮且具有弹性的描述[22],将苓块大,表面黑色,质地坚实,肉质白色的猪苓划为甲级[23];苓种小,表面灰色,苓体烂碎,褶皱不实,肉质褐色者划为乙级[23]。
随着检测技术的发展与应用,评价标准已经转变为以药效成分如甾体化合物和多糖为依据。尽管普遍观点认为猪屎苓的质量优于鸡屎苓[16],目前只有极少数文献对这两种药材进行了明确的化学成分含量检测,且这些检测结果并不能确切地证明两者的品质差异。Zhou[18]检测了12个省区35批猪苓样品中5种甾酮化合物的含量,发现就同一产区而言,猪屎苓和鸡屎苓的甾酮含量并无显著性差别,河南省1个鸡屎苓样品的甾酮总含量高于3个猪屎苓样品的平均值,陕西省2个鸡屎苓样品的甾酮总含量低于9个猪屎苓样品的平均值。Xia等[24]采用HPLC法和水提醇沉法测定了17批猪苓样品的麦角甾醇和多糖含量,结果显示,同一地区猪屎苓的麦角甾醇和多糖含量普遍高于鸡屎苓,但四川省1个野生鸡屎苓样品的多糖含量高于9个栽培猪屎苓的平均含量,所以有学者认为不能仅根据外观形状判断猪苓药材的品质[14]。
陕西地区因其独特的自然条件,被广泛认可为猪苓的优质产区,尤其是太白山区的猪苓,以其优良的品质受到业界的广泛认可[25]。Zhou[26]、Li和Liang[27]分别测得陕西宁强和留坝的猪苓多糖含量最高。Zhou[18]利用HPLC法测定不同猪苓样品中5种甾酮类化合物含量以及对陕西产区的11批猪苓样品的指纹图谱相似度进行分析,结果显示,陕西产区猪苓菌核质量好且稳定性高,这与Zhao等[28]和Li等[29]的实验结果一致。Guo等[30-31]测得陕西猪苓饮片多糖含量最高并且陕西猪苓药材的麦角甾醇、麦角甾-4,6,8(14),22-四烯-3-酮及多糖含量普遍高于其他产地。尽管有个别报道[32]指出陕西地区存在一定量低于药典标准的不合格药材,但这并不影响陕西作为猪苓优质产区的地位。然而,虽然有个别文献[31]表明吉林长春猪苓的总多糖含量高于其他5个产区,目前尚未发现东北地区猪苓在甾体化合物或多糖含量上普遍优于其他地区的科学记载。陕西等地多为猪屎苓、东北等地多为鸡屎苓[19],所以认为猪屎苓的质量优于鸡屎苓的假说具有一定的合理性。然而,鉴于目前文献中对鸡屎苓样品的质量检测相对有限,这一假说的验证仍需更多充分的实验数据和文献支持。
3 猪屎苓和鸡屎苓的遗传差异分析
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种质资源是药材的生产源头,直接关系着药材产量和质量。为了系统地评价猪苓种质资源,指导现存资源的保护和利用,不同学者采用多种方法对猪苓遗传多样性进行了研究。
目前,相关研究表明,不同形态的猪苓是同一个物种。Chen等[33]比较猪屎苓、鸡屎苓和铁蛋屎苓的酯酶(EST)同工酶和过氧化酶(POD)同工酶,发现3种猪苓的EST同工酶带的酶谱型完全相同,而POD酶谱呈现3种谱型。根据POD酶谱相似性指数聚类分析发现,猪屎苓与铁蛋猪屎苓聚成一束,亲缘关系较近,鸡屎苓与其他两种猪苓的亲缘关系相对较远,但三者在酶谱相似度指数大于0.5的范围内仍聚为一类,认为它们属同种。Liu等[9]分别扩增了猪屎苓、鸡屎苓和铁蛋屎苓的rDNA(18rDNA,ITS1-5.8S rDNA-ITS2)和β-tub1序列,扩增出的同源片段碱基序列完全一致,表明菌核形态不同的猪苓为同一物种。
早期研究主要利用同工酶技术对不同形态猪苓进行遗传多样性分析。Xu等[19]对鸡屎苓和猪屎苓进行EST同工酶分析,发现2种菌株没有相对迁移率(Rf值)相同的酶带,认为陕西猪屎苓和长白山区鸡屎苓的遗传差异较大,亲缘关系较远。他经过调研发现长白山区生长的野生猪苓均为鸡屎苓[7],没有猪屎苓,所以提出鸡屎苓是长白山区特异的猪苓种质资源的观点,认为华北、西北地区生长的猪苓多为猪屎苓,东北长白山区生长的多为鸡屎苓[19]。同工酶的表达受生物发育过程中基因的控制,因此同一生物在不同生长阶段表现出的酶谱可能不同[34],这使得利用同工酶技术进行遗传多样性分析具有一定的局限性。
随着分子生物学的发展,分子标记被广泛应用于猪苓遗传多样性、亲缘关系、种群结构的研究。相关报道显示猪苓的亲缘关系与地理分布之间存在一定的关联,东北猪苓与其他地区猪苓亲缘关系较远。Xing等[35]对12个省42个地区猪苓菌核的nrDNA ITS和28S rRNA(LSU)序列进行比对分析并构建了基于ITS+LSU序列的系统发育树,发现全国的猪苓可以分为4个大的类群,其中东北(吉林、辽宁、黑龙江)的鸡屎苓为单独的类群。Yang等[36]基于5个不同省份的7批猪苓的ITS序列构建系统发育树,结果显示,6批样品的系统发育关系紧密聚为一支,而吉林省延边朝鲜族自治州野生猪苓自成一支。
在分子系统发育研究中,样本量的增加对于揭示亲缘关系和地理分布之间的关系至关重要。有学者利用SRAP和EST-SSR引物对不同地区的猪苓菌株进行聚类分析[37-38]。然而,由于这些研究中涉及的样本量较少,尤其是在陕西以外的地区仅包含了单一菌株的样本,因此研究结果可能无法准确反映猪苓亲缘关系与地理分布之间关系。而从Liu等[39]采用ISSR标记对11个不同地区的37份猪苓的分析结果看,东北猪苓与其他地区的区别显著,无论是聚类分析还是主成分分析,吉林及黑龙江的猪苓中均单独归为一类。Liu等[39]进一步基于猪苓转录组数据设计了50对SSR引物对42份猪苓种质进行了遗传多样性分析。研究发现,2个分支的猪苓个体在系统发育树上的聚类与地理位置显著相关,其中一支的猪苓全部来自东北三省,这暗示它们之间亲缘关系相近,而与陕西等地的猪苓亲缘关系较远。同时,研究也发现吉林地区的部分猪苓也与陕西、云南等地区的猪苓聚为一支,这表明随着样品来源的增加,即便在同一地区,也可能存在着遗传上相距甚远的猪苓。Xing等[35]在其研究中发现,来自青海省的猪屎苓样品形成了单独的分枝,与其他猪屎苓样品聚类较远。这说明即使具有相同形态特征的菌核,也可能属于不同的系统发育类型,菌核的形态可能受遗传以外的其他因素影响。
4 猪屎苓和鸡屎苓的菌种栽培潜力
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生产上普遍使用猪苓菌核作为种苓进行无性繁殖生产猪苓,苓种退化使得商品猪苓质量良莠不齐,无节制的乱采滥挖也使得我国野生猪苓资源日益匮乏。猪苓菌种栽培是指将猪苓菌丝形成的菌核作为种苓,与蜜环菌伴栽生产猪苓的方法。采用人工培养的猪苓菌种代替野生菌核人工栽培是解决猪苓种苓紧缺、实现猪苓可持续化生产的有效途径之一。
不少科研工作者先后对猪苓菌丝形成菌核进行了研究。Wang等[40]从野生猪苓菌核中分离到猪苓菌株,发现其在小米培养基上可形成具有黑色外皮的小菌核。Guo等[41]将猪苓菌丝与伴生菌共培养,发现无论是在实验室培养基上还是树棒上,均能发育成正常的猪苓菌核。Wang等[42]在微观水平上证明菌丝形成的菌核和野外获得的菌核结构相似,都有皮层和髓部。Chen等[43]分别采用木屑培养基和木屑棉籽壳混合培养基培养原种和栽培种,发现在适宜的空气湿度和温度下,栽培袋口易形成纯白色且生长较快的菌核组织。目前,关于菌丝形成菌核的机制研究仍处于早期阶段,并且尚未广泛地应用于生产实践中。尽管有极个别文献报道某些菌种栽培技术在小规模的田间试验中取得了一定的成功[44-45],但在实际栽培区域,尚未观察到这些技术成功应用的案例。
要实现猪苓菌种的人工栽培,筛选适宜的猪苓菌株是关键。有研究报道,东北来源的猪苓菌株具有许多优良的培养特性。Liu等[46]从17株不同来源的猪苓菌株中筛选出5个不易老化、多糖含量高的优良菌株,其中2个菌种由东北野生猪苓中分离得到,1个菌种由东北食用菌研究所提供,而Li等[47]从10株猪苓菌株中筛选出的多糖含量和菌丝生长速率最高的菌株也由东北食(药)用真菌研究所提供。鉴于东北地区主要分布着鸡屎苓,我们推测鸡屎苓菌株相较于猪屎苓菌株可能具有更快的生长速度和更高的菌丝多糖含量,这使得它们可能更适合作为菌种进行人工栽培。此外,鸡屎苓菌株可能还具备其他尚未被充分认识的潜在优势。Xu等[19]和Chen等[48]报道鸡屎苓和猪屎苓菌株在生长特性上有明显区别,鸡屎苓菌株生长速度快,不容易老化褐化,能形成菌核状组织。有学者还发现黑龙江来源的猪苓菌丝生长速度最快,最易产生菌核和厚垣孢子,认为其大田繁殖能力和抗逆境能力强,适合用于大田栽培生产[49]。
5 讨论
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本研究从形态、品质、遗传和菌种栽培4个方面对2种不同形态猪苓的相关研究进行了系统梳理和归纳。整理了猪屎苓和鸡屎苓的差异性、探讨了猪苓形态与品质的相关性、分析了猪苓遗传相似性与地理位置的关系以及评估了猪屎苓和鸡屎苓在菌种栽培方面的潜力。总体来看,猪屎苓与鸡屎苓在这4个方面上均存在差异,这些差异可能源于它们的遗传变异以及对不同环境条件的适应。然而,目前的基础研究和应用研究已不能很好地满足猪苓产业高质量发展的需求。因此,未来的研究需要对猪屎苓和鸡屎苓的差异进行更系统、更深入的研究,以便更有效地指导生产实践。
在形态方面,不同研究者对猪苓的描述常带有主观色彩,导致对猪屎苓和鸡屎苓的形态特征描述存在一定的重叠。例如“长形块状”这一术语既被用于描述猪屎苓,也被用于描述鸡屎苓[9,50]。实际上,采集到的猪苓样品在形态上并不总是严格符合传统上对猪屎苓和鸡屎苓的分类标准(图2)。除了常见的猪屎苓和鸡屎苓外,还有如甘肃陇南报道的马屎状猪苓[10],以及秦巴山区的铁蛋苓[8-9],其菌核近似球形或扁块状、基本无分枝、质地较硬。康巴苓[8]菌核有分枝,质地十分疏松,仅在陕西汉中有过报道。这些形态的多样性表明,尽管以菌核大小、形状、表面特性等形态特征来区分猪屎苓与鸡屎苓具有一定的科学依据,但为了提高分类的准确性和客观性,建立更加科学和量化的指标是很必要的。
在品质方面,尽管普遍存在一种观点认为猪屎苓的品质优于鸡屎苓,但目前缺乏足够的科学证据来支持这一假说。现有的文献资料表明,无论是基于甾酮类化合物还是多糖含量的检测,都无法充分证明猪屎苓的品质普遍优于鸡屎苓。研究显示,陕西太白山区的猪苓质量较好[28-31],而较少东北猪苓有效成分含量高于其他地区的记载,陕西多为猪屎苓,东北多为鸡屎苓,所以我们认为猪屎苓的质量优于鸡屎苓的观点具有一定的依据,但缺乏充足的证据予以佐证。遗传背景、栽培方式、地域差异和生长环境等因素都可能影响猪苓的品质,为更科学地评价猪屎苓与鸡屎苓,后续研究应在全国收集猪苓资源,对2种不同形态的猪苓菌核进行全面、系统的评价,以揭示猪苓外观形态与品质之间的相关性。
在遗传方面,目前研究表明,猪屎苓和鸡屎苓属于同一个物种[9,33]。东北猪苓与陕西等地的猪苓不仅形态差距大,亲缘关系也较远。尽管如此,随着样本来源的增加,同一地区内也出现了亲缘关系较远的菌株[39],且相同形态的菌核也可能属于不同的系统发育类型[35]。这些观察结果表明,猪苓菌核表型的分化可能是由遗传、地理、环境或多种因素共同作用的结果。我们在实地考察时还发现,有农户在吉林白山市采挖到野生的鸡屎苓,在田间与蜜环菌伴栽两年后,体形变大,形如陕西的猪屎苓(图3)。这说明猪屎苓和鸡屎苓在外观上的区别可能是由环境因素造成的。猪苓在贫瘠的环境下表面皱缩,为鸡屎苓;在外界营养充分时,体形变大,形成猪屎苓,也有文献[51]报道适宜的生态环境和肥沃的土壤能提高猪苓菌核体积。但目前对猪苓遗传多样性的研究仍然较少,为了更系统地研究猪苓的种质关系等遗传信息,研究猪苓表型分化的影响因素,还需增加样品来源,提高分子标记准确性,开展种系发生、地理起源推断及环境适应性等方面的研究。
菌种栽培技术具有条件可控、易规模化、成本低廉、生态环境友好等优点,是解决猪苓种苓紧缺问题的有效途径之一。如果能够突破该技术,将大大节约种苓和保护野生菌核资源,对猪苓可持续发展、农民增收都具有重大意义。研究表明,鸡屎苓菌株能形成菌核状组织[19],可能更适宜用作菌种栽培猪苓。除此之外,鸡屎苓菌株生长快、抗逆性强的特性可能有望解决目前菌种栽培技术存在的结苓小、不易存活等问题[45]。本课题组及国内其他科研人员长期致力于猪苓菌丝形成菌核的相关研究工作,虽在实验室条件下取得了一些进展[52-54],但目前还未能应用于生产。有研究报道从野生猪苓子实体和菌核中分离的11株猪苓菌株中只有2株可以形成菌核[44],有学者开展了4种菌种的大田栽培试验,发现猪苓菌丝消失,均未采集到菌核,可能是由于菌种抗逆性差,不能适应野生环境[55]。因此,在野外林间进行菌种的实地栽培才是验证菌种是否能用于生产实践的直接证据。
猪苓和蜜环菌属于真菌间较特殊的共生关系[56],猪苓依靠蜜环菌提供营养成分,不同的环境可能会影响蜜环菌种群的组成,从而对当地菌株的遗传结构形成选择压力。Xing等[57-58]基于系统发育网络分析揭示了猪苓-蜜环菌系统发育的相关性,亲缘关系较近的猪苓类群更倾向于与亲缘关系越近的蜜环菌共生,并且蜜环菌在共生关系上表现出更显著的系统发育信号,这都暗示蜜环菌可能是影响猪苓种群结构的因素之一[59]。Li等[29]测定不同产地野生猪苓样品中麦角甾醇含量,发现麦角甾醇含量较高的4个地区的猪苓菌核上长有明显的蜜环菌索,所以推论猪苓菌核的麦角甾醇含量高低可能与其环境中的蜜环菌有关。具体蜜环菌对猪苓质量品质、种群结构的影响还有待进一步的研究。
基金
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  • 中国医学科学院医学与健康科技工程-前沿生物医药技术创新转化研究专项(2022-I2M-2-001)
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2024年第59卷第23期
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doi: 10.11669/cpj.2024.23.001
  • 接收时间：2024-05-17
  • 首发时间：2025-11-13
  • 出版时间：2024-12-08
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  • 收稿日期：2024-05-17
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中国医学科学院医学与健康科技工程-前沿生物医药技术创新转化研究专项(2022-I2M-2-001)
中国医学科学院医学与健康科技创新工程项目(2021-I2M-1-031)
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    中国医学科学院北京协和医学院药用植物研究所, 北京 100193

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* 郭顺星,男,博士,教授 研究方向:药用植物内生真菌生物学 Tel:(010)57833259
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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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