QI Hongzhi,WANG Huifeng,YAN Zhaoling,et al.Research Progress on Growth Conditions and New Cultivation Substrates of Ganoderma[J].Northern Horticulture,2025,(21):122-132.[doi:10.11937/bfyy.20250905]
灵芝生长条件及新型栽培基质研究进展
- Title:
- Research Progress on Growth Conditions and New Cultivation Substrates of Ganoderma
- 文章编号:
- 1001-0009(2025)21-0122-11
- Keywords:
- Ganoderma; biological characteristics; growth conditions; cultivation substrate; active ingredient
- 分类号:
- S 567.3+1
- 文献标志码:
- A
- 摘要:
- 灵芝富含多糖、三萜等多种药用成分，是我国传统中药材，同时也是药食同源类真菌。随着大健康产业的发展，灵芝需求量剧增，对灵芝人工栽培提出了更高的要求。从营养条件（最佳碳源、氮源、无机盐、维生素等）和环境条件（最适温度、湿度、光照、空气、pH等）2个方面分析了不同灵芝菌种所需的生长条件，并从农作物废料、林果木废弃物、中药材废弃物、菌草等方面系统综述了灵芝新型栽培基质方面的研究进展，最后指出今后应从野生灵芝资源保护和筛选、因地制宜开发灵芝新型栽培基质、菌株与基质标准化配套技术、基质和灵芝的食用安全性等方面加强灵芝栽培研究，以期为灵芝人工栽培产业的发展提供参考依据。
- Abstract:
- Ganoderma is rich in polysaccharides，triterpenes and other medicinal ingredients.It is a traditional Chinese herbal medicine in China，and it is also a medicinal and edible fungus.With the development of the health industry，the demand for Ganoderma has increased dramatically，which puts forward higher requirements for the artificial cultivation of Ganoderma.The growth conditions of Ganoderma strains were analyzed from nutritional conditions (optimum carbon source，nitrogen source，inorganic salt，vitamin，et al.) and environmental conditions (optimum temperature，humidity，light，air，pH，et al).The research progress of new cultivation substrates of Ganoderma was systematically reviewed from the aspects of crop waste，forest and fruit wood waste，Chinese herbal medicine waste，and Juncao.Finally，it was pointed out that the research on Ganoderma cultivation should be strengthened from the aspects of protection and screening of wild Ganoderma resources，development of new cultivation substrates of Ganoderma according to local conditions，standardized matching technology of strains and substrates，and edible safety of substrates and Ganoderma，so as to provide reference for the development of artificial cultivation industry of Ganoderma.
参考文献/References:
［1］王恬恬,徐济责,刘晓龙.灵芝栽培研究进展［J］.菌物研究,2024,22(1):94-102.［2］国家卫生健康委、市场监管总局.关于党参等9种新增按照传统既是食品又是中药材的物质公告［EB/OL］.(2023-11-17)［2024-12-25］.http://www.nhc.gov.cn/sps/s7892/202311/f0d6ef3033b54333a882e3d009ff49bf.shtml.［3］张波,叶雷,谭伟,等.莲子壳主料对代栽灵芝生长及其生产效益的影响［J］.菌物学报,2024,43(2):90-101.［4］GOH Y K,MARZUKI N F,TAN S Y,et al.Experimental mixture design as a tool to optimize the growth of various Ganoderma species cultivated on media with different sugars［J］.Mycology,2016,7(1):36-44.［5］HU Y,XU W,HU S,et al.In Ganoderma lucidum,Glsnf1 regulates cellulose degradation by inhibiting GlCreA during the utilization of cellulose［J］.Environmental Microbiology,2020,22(1):107-121.［6］尚陆娥,孙达锋,李建英,等.白肉灵芝液体培养基配方优化及代料栽培技术［J］.中国食用菌,2023,42(3):79-83.［7］罗丽媛,荆杨梦杰,刘欣瑞,等.松杉灵芝的生物学特性及不同栽培模式下的生长及活性成分［J/OL］.菌物研究,(2024-12-23)［2025-02-21］.https://doi.org/10.13341/j.jfr.2023.1703.［8］杨学义,董鹏,张波,等.松杉灵芝菌丝生物学特性及其木质化机理初探［J/OL］.菌物研究,(2024-07-05)［2025-02-21］.https://doi.org/10.13341/j.jfr.2023.1691.［9］袁保京,张日俊.碳氮源对灵芝液体发酵胞外糖肽产量的影响［J］.中国农业大学学报,2012,17(1):119-124.［10］LIU L,FENG J,GAO K,et al.Influence of carbon and nitrogen sources on structural features and immunomodulatory activity of exopolysaccharides from Ganoderma lucidum［J］.Process Biochemistry,2022,119:96-105.［11］杨宗渠,李长看,雷志华,等.以小麦为基质的灵芝固体发酵条件研究［J］.江苏农业科学,2017,45(13):150-152.［12］冯杰,冯娜,贾薇,等.氮源对灵芝菌丝体液态深层发酵合成灵芝三萜的影响［J］.菌物学报,2016,35(6):722-733.［13］NGUYEN L T,LE V V,NGUYEN B T T,et al.Optimization of mycelial growth and cultivation of wild Ganoderma sinense［J］.Biotechnologia,2023,104(1):65-74.［14］王庆武,丛倩倩,谢新国,等.不同碳氮比培养基对泰山赤灵芝农艺性状的影响［J］.中国食用菌,2017,36(2):27-29,32.［15］马博,王凯,周树,等.野生灵芝BSU01的分离鉴定、生物学特性及其木质纤维素酶活分析［J］.基因组学与应用生物学,2019,38(3):1109-1116.［16］颉颖睿.代料栽培灵芝的配方筛选及灵芝代谢组学和转录组学研究［D］.郑州:河南中医药大学,2023.［17］田润,陈丽芳,赵润祥,等.亚弯柄灵芝的生物学特性及其驯化栽培［J］.菌物学报,2024,43(10):48-65.［18］何俊,李娥贤,牛开阳,等.滇中灵芝和可食灵芝生物学特性及驯化［J］.食用菌学报,2023,30(4):21-30.［19］雷萍,张文隽,郝哲,等.野生灵芝分离鉴定、生物学特性及活性成分分析［J］.西北农业学报,2022,31(12):1615-1624.［20］ZHANG B,ZHOU J,LI Q,et al.Manganese affects the growth and metabolism of Ganoderma lucidum based on LC-MS analysis［J］.PeerJ,2019(7):e6846.［21］TAJIK A,SAMADLOUIE H R,SALEK FARROKHI A,et al.Optimization of chemical conditions for metabolites production by Ganoderma lucidum using response surface methodology and investigation of antimicrobial as well as anticancer activities［J］.Frontiers in Microbiology,2024(14):1280405.［22］富振洋,杨璐,孙宁,等.不同营养物质对灵芝菌丝生长的影响［J］.中国食用菌,2024,43(2):82-86.［23］韦会平,刘正宇,谭杨梅,等.温度条件对金佛山灵芝生长的影响［J］.中国农学通报,2005,21(10):85-87.［24］刘欣怡,陈敏,雷雅婷,等.赤芝菌草工厂化栽培环境条件优化［J］.北方园艺,2019(12):122-131.［25］康晟菀,周启武,刘迪,等.可控环境因子对白肉灵芝菌丝生长的影响［J］.北方园艺,2023(23):107-114.［26］陈爽,刘淑娟,高悦,等.野生有柄灵芝生物学特性及驯化栽培［J］.菌物学报,2023,42(11):2218-2230.［27］吕艳聪,李玉,张国亮,等.3株野生赤芝的生物学特性及灵芝酸类成分含量的测定［J/OL］.菌物研究,（2023-10-17）［2025-02-21］.https://doi.org/10.13341/j.jfr.2023.1628.［28］ZHANG B,ZHOU J,LI X,et al.Temperature affects substrate-associated bacterial composition during Ganoderma lucidum hyphal growth［J］.Canadian Journal of Microbiology,2021,67(4):281-289.［29］陈合,陈立,陈明,等.中药渣用于培养灵芝菌及功能成分分析［J］.农业工程学报,2006,22(S2):167-170.［30］石云龙,韩梅,孙卓,等.林下栽培灵芝主要活性成分与生态因子的关系［J］.吉林农业大学学报,2024,46(2):239-245.［31］张鑫,邬柯奕,祝嗣臣,等.栽培条件对松杉灵芝多糖含量的影响及其抗氧化活性分析［J］.生物技术通报,2024,40(11):47-57.［32］ZHANG W X,TANG Y J,ZHONG J J.Impact of oxygen level in gaseous phase on gene transcription and ganoderic acid biosynthesis in liquid static cultures of Ganoderma lucidum［J］.Bioprocess and Biosystems Engineering,2010,33(6):683-690.［33］姚珂,韩嘉钰,贺黎铭,等.空气中CO2浓度对灵芝三萜的影响及其分子机制研究［J］.中国食用菌,2018,37(6):44-49.［34］金鑫,张璐,吴鹏,等.遮光处理对盆景灵芝生长发育和酶活性的影响［J］.中国农业科技导报,2023,25(4):147-156.［35］袁学军,陈永敢,陈光宙,等.不同光照和栽培基质对灵芝活性成分的影响［J］.中国食用菌,2012,31(6):38-39,43.［36］颉颖睿,陈向东,仇冠盛,等.不同栽培条件对灵芝质量的影响［J］.时珍国医国药,2023,34(7):1738-1742.［37］梅锡玲,陈若芸,李保明,等.光质对灵芝菌丝体生长及三萜酸量影响的研究［J］.中草药,2013,44(24):3546-3550.［38］梅锡玲,赵洲,陈向东,等.光质对灵芝菌丝体生长及内源IAA代谢调控的研究［J］.中国中药杂志,2013,38(12):1887-1892.［39］ROSALES-LPEZ C,ARCE-TORRES F,MONGE-ARTAVIA M,et al.Evaluation of the use of elicitors for the production of antioxidant compounds in liquid cultures of Ganoderma curtisii from costa rica［J］.Molecules,2022,27(13):4265.［40］LIU D,SUN X,DIAO W,et al.Comparative transcriptome analysis revealed candidate genes involved in fruiting body development and sporulation in Ganoderma lucidum［J］.Archives of Microbiology,2022,204(8):514.［41］刘凌云,黄在兴,邢世和,等.灵芝生长过程中培养料中的碳转化及CO2排放［J］.园艺学报,2019,46(10):2047-2054.［42］朱宗财,高能,孔新,等.棉花废弃物栽培食用菌基质研究进展［J］.北方园艺,2024(19):126-132.［43］曹正,谢春芹,凡军民,等.茶园套种灵芝仿生栽培模式的研究［J］.中国食用菌,2018,37(4):26-29.［44］王爱仙,王怡暄,张闽春,等.不同栽培配方对灵芝产量与品质的影响［J］.中国食用菌,2021,40(2):23-27.［45］陈宜,谢晟,李胜杰,等.棉花秸秆栽培灵芝的品种与配方筛选试验［J］.中国食用菌,2016,35(4):47-49,53.［46］冀宏,王谦,汪虹,等.玉米秸袋栽金针菇灵芝科学依据初探［J］.中国食用菌,2001,20(6):10-12.［47］陈永敢,范平杰,李珍,等.玉米秆在灵芝制种中的应用［J］.北方园艺,2016(12):144-146.［48］李林,郭红艳,孙晓杰,等.利用大豆秸杆和玉米秸秆栽培灵芝［J］.食用菌学报,2021,28(4):15-19.［49］魏雅冬,李艳芳,戴明,等.玉米芯和玉米秸栽培灵芝、猴头及榆黄蘑配方研究［J］.陕西农业科学,2017,63(7):12-14.［50］徐恒,董爱荣,申东辰,等.树舌灵芝栽培料配方比较试验［J］.食用菌,2024,46(6):29-31.［51］花纪,颜俊清,霍光华,等.无柄灵芝代料栽培基质营养组成与其功能成分累积的关系研究［J］.食品科技,2019,44(6):38-44.［52］赵霞,王会锋,杨豫龙,等.药用灵芝育种栽培及品质研究进展［J］.乡村科技，2024，15(16)：86-90.［53］韩嘉钰,魏巍,余梦瑶,等.不同栽培基质对灵芝基质营养物质及胞外酶活的影响［J］.时珍国医国药,2016,27(3):715-719.［54］MANAVALAN T,MANAVALAN A,THANGAVELU K P,et al.Secretome analysis of Ganoderma lucidum cultivated in sugarcane bagasse［J］.Journal of Proteomics,2012,77:298-309.［55］赵风云,杨鑫,黄淑君,等.以蔗渣和棉籽壳为基质的灵芝酶活性及其营养对比研究［J］.现代食品科技,2015,31(1):84-89.［56］黄清铧,王庆福,高裕锋,等.甘蔗渣、啤酒糟培养灵芝研究和成分分析［J］.食品工业科技,2015,36(21):143-147.［57］VEENA S S,PANDEY M.Paddy straw as a substrate for the cultivation of Lingzhi or Reishi medicinal mushroom,Ganoderma lucidum (W.Curt.Fr.) P.Karst.in India［J］.International Journal of Medicinal Mushrooms,2011,13(4):397-400.［58］C〖JX-+0.8mm〗〖KG-*2〗＇ILERDIC〖JX-+0.8mm〗〖KG-*2〗＇J,VUKOJEVIC〖JX-+0.8mm〗〖KG-*2〗＇J,STAJIC〖JX-+0.8mm〗〖KG-*2〗＇M,et al.Biological activity of Ganoderma lucidum basidiocarps cultivated on alternative and commercial substrate［J］.Journal of Ethnopharmacology,2014,155(1):312-319.［59］NEDELJKOVIC B B,C〖JX-+0.8mm〗〖KG-*2〗＇ILERDIC〖JX-+0.8mm〗〖KG-*2〗＇J,ZMIJANJAC D,et al.Immunomodulatory effects of extract of Lingzhi or reishi medicinal mushroom Ganoderma lucidum (Agaricomycetes) basidiocarps cultivated on alternative substrate［J］.International Journal of Medicinal Mushrooms,2022,24(8):45-59.［60］陈永敢,屈直,李珍,等.花生壳在灵芝制种中的应用研究［J］.种子,2016,35(5):129-131.［61］BIDEGAIN M A,CUBITTO M A,CURVETTO N R.Optimization of the yield of Lingzhi or reishi medicinal mushroom,Ganoderma lucidum (higher basidiomycetes),cultivated on a sunflower seed hull substrate produced in Argentina:effect of olive oil and copper［J］.International Journal of Medicinal Mushrooms,2015,17(11):1095-1105.［62］RASHAD F M,EL KATTAN M H,FATHY H M,et al.Recycling of agro-wastes for Ganoderma lucidum mushroom production and Ganoderma post mushroom substrate as soil amendment［J］.Waste Management,2019,88:147-159.［63］覃炎锋,李荷,巩伯梁,等.响应面法优化以桑枝为主要基质的灵芝栽培配方［J］.微生物学免疫学进展,2016,44(3):36-42.［64］舒庭辉.不同栽培原料对灵芝生长及子实体营养品质的影响［J］.特种经济动植物,2023,26(12):39-41.［65］张波,谭伟,周洁,等.不同颗粒度桑枝屑代料栽培灵芝产量和品质差异［J］.中国农学通报,2022,38(22):39-43.［66］贺望兴,石旭平,胡佳,等.茶枝屑代料栽培灵芝配方优选［J］.安徽农业科学,2019,47(6):52-54,58.［67］刘明香,林忠宁,陈敏健,等.茶枝屑代料栽培灵芝菌株的筛选［J］.福建农林大学学报(自然科学版),2012,41(2):183-186.［68］刘新华,吕军,董学堂,等.以果树枝木屑为主料栽培灵芝及其菌渣栽培平菇试验［J］.食用菌,2017,39(4):31-33.［69］LUO G,PAN Z,LIU Z,et al.Influence of cultivation substrate on antioxidant activities and triterpenoid profiles of the fruiting body of Ganoderma lucidum［J］.Frontiers in Nutrition,2024,11:1329579.［70］吕成杰,孙莹,刘伟,等.用苹果树枝栽培灵芝试验［J］.中国食用菌,2016,35(6):86-88.［71］赵威,赵传岳,周婷,等.“灵芝+白星花金龟幼虫” 联合转化苹果修剪枝条研究［J］.山东农业大学学报(自然科学版),2023,54(5):710-717.［72］杨仁德,陈萍,杨杰仲,等.猕猴桃树枝栽培灵芝基质配方筛选试验［J］.现代化农业,2018(9):33-34.［73］MORALES-ESTRADA I,GAITN-HERNNDEZ R,GUTIRREZ A,et al.Vineyard pruning waste improves bioconversion and chemical composition of native Ganoderma spp.(Agaricomycetes) strains from Mexico［J］.International Journal of Medicinal Mushrooms,2018,20(8):775-789.［74］丁李春.废弃葡萄枝蔓栽培灵芝的适宜菌株筛选［J］.食药用菌,2021,29(2):148-151.［75］赖腾强,谢娜,吴伯文,等.不同枇杷枝屑配方对灵芝栽培特性和品质的影响分析［J］.农业与技术,2022,42(3):15-17.［76］张平,谢娜,赖腾强,等.枇杷枝屑代料栽培对不同菌株灵芝品质的影响［J］.北方园艺,2017(14):165-168.［77］高静静,庄新辉,谭雪松,等.中药渣资源化利用研究进展［J］.中成药,2024,46(2):536-541.［78］霍光明,张李阳,张吉.不同栽培基质对灵芝品质的影响［J］.南京晓庄学院学报,2017,33(6):92-94.［79］蔡良,欧阳冬青,施华蓉,等.不同中药配方颗粒药渣栽培灵芝研究［J］.福建农业科技,2020,51(9):39-44.［80］张鑫,谢苗,亓小妮,等.适宜药渣栽培的灵芝优良菌株筛选与鉴定［J］.中国食用菌,2021,40(9):34-39.［81］夏伯阳,夏振华,庞盼盼,等.中药渣栽培灵芝的基质配方研究［J］.农村经济与科技,2021,32(8):28-30.［82］刘代缓,韩颜超,时登龙,等.3种中药渣栽培5种食药用菌利用潜力研究［J］.时珍国医国药,2023,34(10):2483-2485.［83］周晶,林兴生,林辉,等.菌草研究与应用进展［J］.福建农林大学学报(自然科学版),2020,49(2):145-152.［84］ZHENG H,WANG B,HUA X,et al.A near-complete genome assembly of the allotetrapolyploid Cenchrus fungigraminus (JUJUNCAO) provides insights into its evolution and C4 photosynthesis［J］.Plant Communications,2023,4(5):100633.［85］林冬梅,罗虹建,林树钱,等.不同菌草基质栽培灵芝多指标质量评价［J］.福建农业学报,2022,37(12):1579-1585.［86］吴鸿雪,唐庆九,吕旭聪,等.栽培条件对灵芝子实体的三萜组成及其活性的影响［J］.菌物学报,2020,39(1):144-154.［87］吴鸿雪,吕旭聪,唐庆九,等.栽培条件对灵芝子实体中四种重金属含量的影响［J］.菌物学报,2020,39(1):137-143.［88］杨丽秋,范锦琳,刘欣怡,等.菌草对灵芝生长状况及营养成分的影响［J］.福建农业学报,2017,32(5):508-511.［89］刘艳玲,刘朋虎,李晶,等.不同栽培原料对灵芝生长及其子实体营养品质的影响［J］.福建农业学报,2022,37(5):609-616.［90］LIU D,DIAO W,SUN X,et al.Application of Miscanthus substrates in the cultivation of Ganoderma lingzhi［J］.Archives of Microbiology,2023,205(12):384.［91］张卓亿,谢晓方,王霞,等.芦苇、南荻在食用菌栽培中的基质化利用研究进展［J］.北方园艺,2024(24):113-121.［92］林应兴.菌草灵芝研究现状与前景［J］.北方园艺,2021(11):142-147.［93］林兴生,林占熺,林辉,等.菌草鹿角灵芝新菌株栽培特性及活性成分分析［J］.四川大学学报(自然科学版),2020,57(1):181-185.［94］周威杰,罗培娟,成纪予.利用竹屑代料栽培灵芝的试验［J］.浙江农业科学,2022,63(7):1469-1471,1474.［95］付学艳,杜青.利用玉米糖渣栽培灵芝技术要点［J］.食用菌,2018,40(1):57-58.［96］马红梅,陈永敢,袁学军.灵芝的椰子果皮代料栽培试验［J］.江苏农业科学,2013,41(2):204-205.［97］LIU Y,QIAN Y,WANG C,et al.Study of the metabolite changes in Ganoderma lucidum under pineapple leaf residue stress via LC-MS/MS coupled with a non-targeted metabolomics approach［J］.Metabolites,2023,13(4):487.［98］解凡,叶丽云,赵丽丽,等.三株野生重伞灵芝的鉴定及其多糖、三萜含量分析［J］.食用菌学报,2018,25(3):8-14,107.［99］范蕾,刘敏,余乐,等.ICP-MS分析丽水地产灵芝中的重金属元素［J］.中国现代应用药学,2019,36(7):837-840.
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备注/Memo
第一作者简介：齐红志（1985-），女，硕士，助理研究员，现主要从事药用作物栽培等研究工作。E-mail:1240564836@qq.com.责任作者：赵霞（1973-），女，博士，研究员，现主要从事中药材精准生产等研究工作。E-mail:zhaoxia1007@126.com.基金项目：河南省农业科学院优质中药材专项资助项目（ZX20241004）；河南省农业科学院自主创新资助项目（2024ZC070）；河南省特聘研究员资助项目（海外2025）。收稿日期：2025-03-12