药用真菌桑黄的研究进展

桑黄是著名的药用真菌,在抗肿瘤、抗氧化、降血糖、调节免疫等方面有显著功效。目前世界范围内桑黄类群共包括12个种,其中中国分布7个种。本文综述了桑黄物种多样性、形态特征、地理分布、药用成分、药理作用机制以及人工培养等方面的研究进展。     

“桑黄”的本草补充考证

对古代本草著作中记载的"桑黄"的名称、形态特征和药用功效等方面做了补充考证。根据历代本草文献中记载的"桑黄"的特点,将其与现代药用真菌粗毛纤孔菌进行了生境、形态特征和药用功效等3个方面的比较考证,并结合粗毛纤孔菌的现代药理活性方面研究成果,认为古代中药"桑黄"应为多孔菌科真菌粗毛纤孔菌。     

桑黄孔菌属的化学成分及药理作用研究进展

对桑黄孔菌属的分类地位进行梳理和明确,并就桑黄孔菌属内物种的化学成分种类及药理作用的研究进展进行归纳总结。以“桑黄、化学成分、活性、药理作用”等为关键词,检索1960—2020年Web of Science、PubMed、中国知网和中国万方等数据库中桑黄的相关文献,总结了桑黄的化学成分及药理活性研究进展。检索筛选中文文献240篇,外文文献102篇,发现桑黄中含有的化学成分种类及含量丰富,药理作用广泛,包括抗肿瘤作用、抗氧化作用、抗病毒、增强免疫功能、降血糖和降尿酸作用等。通过对中药桑黄的化学成分及其药理作用的归类整理,为深入研究桑黄及其配伍应用、开拓药用范围并明确药效机制等研究打开新思路。     

桑黄类真菌多糖研究进展

桑黄是一类珍稀药用真菌的统称,多糖作为其主要的有效成分之一,具有抑肿瘤、抗氧化、抑菌、消炎、免疫调节等多种药理功效。本文就桑黄多糖的提取分离纯化、结构解析和药用价值等方面的研究进展进行了综述,旨在为深入研究其药理功效并开发成为天然食品、保健品或药品提供重要参考。     

药用真菌桑黄的研究进展

以近年来国内外发表的文献为依据 ,从形态特征、生物学生长特性、主要化学成分、药理学研究等方面综述了文献中称为“桑黄”[包括 :鲍氏针层孔菌 (Phellinusbaumii)、火木针层孔菌 (P .igniarius)、裂蹄针层孔菌 (P .linteus) 3种 ]的药用菌子实体、发酵菌丝体、发酵胞外物的研究进展。结果 :桑黄的人工栽培 (日本、韩国学者采用的室外荫棚段木埋畦栽培法 ) ,固体、液体发酵培养均已获成功。其主要化学成分是子实体、菌丝体多糖以及发酵液中的胞外多糖 ,此外还有黄酮及其衍生物、香豆素类、甾醇类化合物。其子实体、菌丝体及提取物 (主要为多糖 )、胞外多糖在抗发炎、抗氧化、抑制肿瘤生成、增强免疫力、保护肝脏、预防和治疗关节炎等方面有显著的功效。桑黄的各种产品 (包括 :子实体、菌丝体微粉末、提取物浸膏、桑黄茶、桑黄口服液等 )市场需求量很大 ,表明桑黄有广阔的发展前景。     

“桑黄”的本草考证

对“桑黄”的古代、现代名称及其药用功效等方面做了全面考证.“桑黄”最早在汉代的《神农本草经》中记载为“桑耳”,在唐代甄权的《药性论》中才首次出现“桑黄”两字.认为中国“桑黄”,在分类上属于真菌界Fungi、担子菌门Basidiomycota、绣革孔菌目Hymenochaetales、绣革孔菌科Hymenochaetaceae,主要包括来自Phellinus、Inonotus及Fomitiporia等属的具有显著抗肿瘤活性的木腐菌子实体,包括火木层孔菌Phellinus igniarius、瓦尼木层孔菌Phellinus vaninii、鲍姆木层孔菌Phellinus baumii、椭圆嗜蓝孢孔菌Fomitiporia ellipsoidea及粗毛纤孔菌Inonotus hispidus等的子实体,并具有明显的地域特色.     

桑黄菌液体培养过程中酶活及多糖含量变化规律

桑黄孔菌属Sanghuangporus是一类具有重要药用价值的大型真菌,目前被国际公认为抗肿瘤效果最好的真菌之一。本研究以桑黄孔菌属中的杨树桑黄Sanghuangporus vaninii、鲍姆桑黄Sanghuangporus baumii和桑树桑黄Sanghuangporus sanghuang为研究对象,通过测定它们液体培养过程中第3、6、9、12、15、18和21天的菌丝生物量以及发酵液的多糖含量、蛋白质含量、漆酶活性、羧甲基纤维素酶活性、半纤维素酶活性、淀粉酶活性和蛋白酶活性等7个指标,对桑黄的生长代谢能力进行了评价。结果显示,3种桑黄真菌的发酵液均具有完整的胞外酶体系。相比之下,鲍姆桑黄的羧甲基纤维素酶、半纤维素酶和淀粉酶活性更高,杨树桑黄的漆酶活性更高,而桑树桑黄的蛋白酶活性更高,Pearson分析发现多糖的累积与其分泌的羧甲基纤维素酶、淀粉酶和半纤维素酶呈现显著正相关;蛋白质含量则与淀粉酶显著正相关。相应的,鲍姆桑黄和桑树桑黄在多糖含量和蛋白质含量方面显著优于杨树桑黄。研究结果为更好地研究、开发和利用桑黄提供了科学参考。     

杨树桑黄饮片水提物对免疫低下小鼠的免疫功能的影响

探究桑黄饮片水提物对环磷酰胺所致免疫低下小鼠的免疫功能的影响。本研究以桑黄饮片为原料,采用超声和沸水提取方法得到桑黄饮片水提物,注射环磷酰胺造成小鼠免疫功能低下,通过测定脾脏和胸腺指数、血清免疫球蛋白和细胞因子、脾淋巴细胞增殖、碳粒廓清等实验,对免疫低下小鼠的免疫功能进行研究评价。结果表明,灌胃桑黄饮片水提物的小鼠与模型组相比,免疫器官指数、血清中免疫球蛋白含量、细胞因子水平、脾淋巴细胞增殖程度、巨噬细胞的吞噬速率和吞噬指数、小鼠耳肿胀度均有提高,脾脏苏木精-伊红(HE)染色也表明桑黄饮片水提物可以改善环磷酰胺导致的脾脏组织病理状态。本研究表明桑黄饮片水提物对环磷酰胺所致的免疫抑制小鼠具有良好的免疫保护作用,为桑黄饮片扩大临床应用提供实验依据。     

桑黄多糖的药理作用及提取方法研究进展

桑黄是我国珍贵的药用真菌,其药效成分有多糖类、黄酮类、三萜类、甾类、多酚类、吡喃酮类及生物碱等。其中多糖作为桑黄的重要有效成分,具有抗肿瘤、抗氧化、免疫调节、抗疲劳、护肝、降血糖及抗菌抗炎等广泛的药理作用,且无毒副作用,因此具有重要的研究价值。近些年,许多学者对桑黄多糖提取进行了大量的实验研究,结果表明溶剂浸提法、超声波提取法和盐提取法适用于批量工业生产,但溶剂浸提法资源耗费较大;相比而言,微波提取法和酶提取法更适合少量提取用于试验研究,其优点在于提取率高、操作便捷及杂质较少。结合国内外相关研究内容与成果,对桑黄多糖的药理作用及多糖提取技术进行归纳与总结,并对其发展前景进行了展望,旨为桑黄多糖的深入研究提供参考。     

桑黄纤孔菌子实体粉的毒理研究

本文主要研究桑黄纤孔菌子实体粉的毒理学并评价其安全性。利用小鼠急性毒性试验和遗传毒性试验(Ames试验、小鼠骨髓细胞微核试验、小鼠精子畸形试验)对桑黄纤孔菌子实体粉的毒性进行考察。试验结果显示,小鼠对桑黄纤孔菌子实体粉最大耐受量大于12g/kg;桑黄纤孔菌子实体粉在有或无S9的条件下,对组氨酸营养缺陷型鼠伤寒沙门氏菌TA97a、TA98、TA100及TA102均无潜在致突变性;桑黄纤孔菌子实体粉在小鼠骨髓细胞微核试验和小鼠精子畸形试验中均呈阴性反应,不同剂量的样品组与阴性对照组之间无显著性差异。该结果表明桑黄纤孔菌子实体粉基本无毒性,属实际无毒物质。本研究为桑黄纤孔菌子实体粉的产品开发和应用提供了科学依据。     

接种丛枝菌根真菌对枳吸收利用磷酸铝的影响

在温室沙培条件下,以枳实生苗为试材,研究了丛枝菌根真菌摩西球囊霉和地表球囊霉对枳吸收利用难溶性磷酸盐(Al-P)的影响.结果表明：接种菌根真菌显著增加了枳的干物质量、含磷量及磷吸收量,且随Al-P施用量的提高,菌根贡献率、全株磷吸收量、真菌磷吸收量及真菌磷吸收贡献率显著增加;接种处理显著增加了枳根系与菌丝磷酸酶的分泌量,特别是酸性磷酸酶和中性磷酸酶的分泌量,但二者随施磷量的增加而降低.真菌磷吸收贡献率与酸性磷酸酶、中性磷酸酶、碱性磷酸酶和总磷酸酶含量呈极显著正相关.     

Effects of simultaneous infections of endophytic fungi and arbuscular mycorrhizal fungi on the growth of their shared host grass Achnatherum sibiricum under varying N and P supply

Grasses can be infected by endophytic fungi and arbuscular mycorrhizal (AM) fungi simultaneously. Here, we investigated the interactions of a native grass, Achnatherum sibiricum, with both endophytic and AM fungi (Glomus mosseae, GM and Glomus etunicatum, GE) at different nitrogen (N) and phosphorus (P) levels. The results showed that endophyte infection significantly suppressed the colonization rates and spore density of GE, but had no effect on those of GM. Endophyte infection increased shoot biomass regardless of the nutrient conditions. The effects of AM fungi on host growth were dependent on mycorrhizal species. There was no significant interaction between endophytic fungi and GE on host growth; however, a significant interaction between endophytic fungi and GE occurred in total phenolic concentrations and P uptake. As for GM, a significant interaction among endophytic fungi, AM fungi and nutrient availability occurred in shoot growth. Under sufficient N and P conditions, endophyte infection alleviated the detrimental effects of GM colonization on host growth.     

Mutualism is not restricted to tree-killing bark beetles and fungi: the ecological stoichiometry of secondary bark beetles,fungi, and a scavenger

1. All bark beetles are in symbiosis with fungi. Although obligate mutualisms with fungi are common with tree-killing bark beetles (primaries), fungi associated with non-tree-killing bark beetles (secondaries) are usually dismissed as commensals. 2. Using an ecological stoichiometric approach, we show secondaries are also involved in nutrition-based mutualisms, some of which appear obligate, and that differences in symbiont provisioning efficiency have a potent effect on beetle carbon (C): nitrogen (N): phosphorus (P) ratios. 3. Some secondary beetles have high P contents and require efficient P provisioning via fungi, while others have low P contents that may allow them to exploit less efficient fungi or a broader range of species with variable efficiencies. A co-occurring scavenger that feeds on nutrient-poor bark beetle frass (excrement/boring residues) exhibited the lowest phosphorus content yet recorded for an invertebrate. 4. Our results generally support the growth-rate hypothesis that posits differences in C:P and N:P ratios in consumers are due to differential allocation of P to P-rich RNA to support growth. However, while the beetle species that accumulated the most biomass was considerably enriched in P and that with the least biomass was P-poor, one beetle species that was P-rich was also small possibly due to limitation by an element other than P. 5. Our results indicate that fungi are important to a broader range of bark beetles than previously recognised. Additional research is needed to describe how these various symbioses influence forest ecosystems via differential effects of fungi on host beetle fitness.     

植物磷营养状况对丛枝菌根真菌生长及代谢活性的调控*

采用四室隔网培养装置,以玉米为宿主植物,通过在植物生长室设0、50、250和500 mgPkg-1 4个施磷水平,研究了植株体内的磷营养状况对AM真菌Glomus sinuosum和Glomus intraradices生长及活性的影响。研究发现在不施磷条件下,接种AM真菌G. intraradices显著促进了植物生长和磷的吸收;低磷条件(50 mgPkg-1)下,接种菌根真菌显著促进了植物对磷的吸收,但对植物生长没有明显的影响;而在高磷条件(250 mgPkg-1 和500 mgPkg-1)下,接种菌根真菌不但没有促进植物的生长和磷的吸收,反而对其有抑制作用。随着施磷水平的提高, AM真菌根内菌丝的碱性磷酸酶活性显著下降;与不施磷相比,低量(50 mgPkg-1)供磷增加了AM真菌土壤中根外菌丝的密度,高磷(250 mgPkg-1 和500 mgPkg-1)降低了土壤中根外菌丝的密度。上述结果说明:⑴ 给宿主植物施用磷肥引起的植物磷营养状况的改变,对AM真菌生长和代谢活性具有一定的调控作用;⑵ G. sinuosum和G. intraradices两种AM真菌的生长和代谢活性对施磷水平的响应程度无显著性差异;⑶ 高磷抑制AM真菌生长和代谢活性,使真菌吸磷量减少,可能是造成菌根效应降低的原因之一     

Fresh perspectives on the roles of arbuscular mycorrhizal fungi in plant nutrition and growth

Recent research on arbuscular mycorrhizas has demonstrated that AM fungi play a significant role in plant phosphorus (P) uptake, regardless of whether the plant responds positively to colonization in terms of growth or P content. Here we focus particularly on implications of this finding for consideration of the balance between organic carbon (C) use by the fungi and P delivery (i.e. the C-P trade between the symbionts). Positive growth responses to arbuscular mycorrhizal (AM) colonization are attributed frequently to increased P uptake via the fungus, which results in relief of P deficiency and increased growth. Zero AM responses, compared with non-mycorrhizal (NM) plants, have conventionally been attributed to failure of the fungi to deliver P to the plants. Negative responses, combined with excessive C use, have been attributed to this failure. The fungi were viewed as parasites. Demonstration that the AM pathway of P uptake operates in such plants indicates that direct P uptake by the roots is reduced and that the fungi are not parasites but mutualists because they deliver P as well as using C. We suggest that poor plant growth is the result of P deficiency because AM fungi lower the amount of P taken up directly by roots but the AM uptake of P does compensate for the reduction. The implications of interplay between direct root uptake and AM fungal uptake of P also include increased tolerance of AM plants to toxins such as arsenate and increased success when competing with NM plants. Finally we discuss the new information on C-P trade in the context of control of the symbiosis by the fungus or the plant, including new information (from NM plants) on sugar transport and on the role of sucrose in the signaling network involved in responses of plants to P deprivation.     

Phosphorus uptake by a community of arbuscular mycorrhizal fungi in jarrah forest

Communities of indigenous arbusuclar mycorrhizal (AM) fungi are expected to alter phosphorus uptake and biomass productivity of plants according to characteristics of the life cycles of the fungi present and the way they interact with each other inside roots and with host plants. Differences in the relative abundance of AM fungi inside roots could influence P uptake if the fungi present differ in effectiveness at accessing P and transferring it to the plant. However, it is difficult to assess the contribution of AM fungi under field conditions. We investigated P uptake, from point sources of P placed 2, 4 and 6 cm from roots, by plants colonised by a community of AM fungi in jarrah forest soil. Roots were retained within a mesh bag to prevent them from growing towards the point source of P. The relative abundance of morphotypes of fungi inside roots and the P status of plants were assessed after 12 and 16 weeks. First, a bioassay was carried out in undisturbed forest soil cores using two host plants, a forest understorey plant Phyllanthus calycinus Labill and the annual pasture species subterranean clover (Trifolium subterraneaumL.), to assess the infectivity of the indigenous community of AM fungi. Roots of both bioassay host plants were colonised in similar proportions by morphotypes of AM fungi resembling Glomus, Acaulospora, Scutellospora and fine endophytes. In this bioassay, there were positive correlations between the proportion of root length colonised and plant biomass and P uptake for P. calycinus, but not for subterranean clover. In the experiment assessing the capacity of P. calycinus to access P placed at increasing distances from the root, shoot P content and concentration in P. calycinus were greater when P was placed 2 cm compared with 4 and 6 cm from roots. The length of hyphae in the vicinity of the point source of P decreased with increasing distance from the plant. The extent to which the individual AM fungi were involved in P uptake is not known. The Glomus morphotype was dominant at both times of sampling.     

植酸钠对菌根真菌根内菌丝碱性磷酸酶活性及根外菌丝生长的影响

采用三室隔网培养装置,以玉米为宿主植物,接种丛枝菌根真菌(AM)(Glomus intraradices),研究了不同用量的植酸钠对AM真菌生长和代谢活性的影响．研究发现,接种AM真菌的植株地上部和根系的P浓度和吸P量,比非菌根植物的提高了1～2倍．外源植酸钠的存在,显著降低了AM真菌根内菌丝的碱性磷酸酶活性,增加了AM真菌在土壤中的菌丝密度．结果表明,外源植酸钠对根内AM真菌碱性磷酸酶活性和真菌根外菌丝的生长具有调控(增减)作用,并且AM真菌提高了植物对土壤固有养分和外源植酸钠中P的吸收和利用．     

Effects of co-inoculation with arbuscular mycorrhizal fungi and rhizobia on soybean growth as related to root architecture and availability of N and P

Soybean plants can form tripartite symbiotic associations with rhizobia and arbuscular mycorrhizal (AM) fungi, but little is known about effects of co-inoculation with rhizobia and AM fungi on plant growth, or their relationships to root architecture as well as nitrogen (N) and phosphorus (P) availability. In the present study, two soybean genotypes contrasting in root architecture were grown in a field experiment to evaluate relationships among soybean root architecture, AMF colonization, and nodulation under natural conditions. Additionally, a soil pot experiment in greenhouse was conducted to investigate the effects of co-inoculation with rhizobia and AM fungi on soybean growth, and uptake of N and P. Our results indicated that there was a complementary relationship between root architecture and AMF colonization in the field. The deep root soybean genotype had greater AMF colonization at low P, but better nodulation with high P supply than the shallow root genotype. A synergistic relationship dependent on N and P status exists between rhizobia and AM fungi on soybean growth. Co-inoculation with rhizobia and AM fungi significantly increased soybean growth under low P and/or low N conditions as indicated by increased shoot dry weight, along with plant N and P content. There were no significant effects of inoculation under adequate N and P conditions. Furthermore, the effects of co-inoculation were related to root architecture. The deep root genotype, HN112, benefited more from co-inoculation than the shallow root genotype, HN89. Our results elucidate new insights into the relationship between rhizobia, AM fungi, and plant growth under limitation of multiple nutrients, and thereby provides a theoretical basis for application of co-inoculation in field-grown soybean.     

Interaction between Endophytic Fungi and Seedlings of Two Species of Paphiopedilum during Symbiotic Culture

Two fungi PM 1 and PA 1 were isolated from P.micranthum and P.armeniacum, respectively, the former belonged to Epulorhiza while the latter belonged to Tulasnella calospora. In this study, seedlings of P.armeniacum and P.micranthum were inoculated with PM 1 on DE medium, the biomass and the content of Ca, Mn of both P.armeniacum and P.micranthum seedlings were higher than those in the control seedlings, which indicated that PM 1 could improve the growth of those species. Seedlings of P.armeniacum and P.micranthum were inoculated with PA 1 on DE medium, the biomass in both species was no difference with those in the control ones. But PA 1 enhanced the dry weight of P.armeniacum according to the rate of dry weight increase. The impact of PM 1 on P.micranthum was higher than on P.armeniacum, while PA 1 impacted conversely, which demonstrated that fungi had stronger improving effect only on its own host plants instead of any other orchids. P.armeniacum seedlings wereinoculated with PM 1 and PA 1 on Harvais medium. The growth of seedlings was inhibited by fungi. The results indicated that nutrient level of medium played an important role in affecting the symbiotic relationship between P.armeniacum seedings and endophytic fungi. Poor nutrient medium was beneficial to the symbiotic relationship and nutrient rich medium was not conducive to the symbiosis.     

不同水肥条件下AM真菌对丹参幼苗生长和营养成分的影响

利用盆栽接种试验,探讨不同水肥条件下AM真菌摩西球囊霉Glomus mosseae对丹参幼苗生长和微量元素的影响,为丹参水肥合理施用提供理论依据。结果表明,不同水肥条件下,接种AM真菌显著提高了根系侵染率和生物量。40%相对含水量、不同施P水平,接种株丹参酮含量升高,总黄酮、丹参素及地下部总酚酸含量降低,植株Zn及地上部Ca、K、Mn、Fe含量升高,而对植株Mg、Cu和地下部Ca、K、Mn、Fe无显著影响;接种效应随施P量不同而变化。70%相对含水量、不同施P水平,接种株药用成分含量显著升高,植株Ca、Mn和地上部K、Cu及地下部Fe和Zn含量升高,而对植株Mg、地下部K、Cu和地上部Fe和Zn含量无显著影响。不同水分和同一施P水平,接种株丹参酮含量升高,地上部Ca、K和地下部Zn含量升高,接种效应因土壤含水量不同而变化,其中以70%含水量时效果最好。说明AM真菌能促进宿主植物根系对水分和矿质元素的吸收与利用,提高水分和P肥利用率,降低水分和P胁迫对丹参的伤害程度,其中以70%相对含水量,施P量为0.16 gP/kg土时AM真菌对丹参的接种效果最佳。