双孢蘑菇覆土出菇机理研究进展

doi:10.16420/j.issn.0513-353x.2022-0460

摘要/Abstract

摘要：

双孢蘑菇栽培必须覆土才能出菇，限制了其产业的高效发展。本文中综述了乙烯和1-辛烯-3-醇对双孢蘑菇覆土出菇影响的机理研究进展，包括抑制出菇的气体自抑物质的合成和作用机制等，并对未来的研究方向进行了展望。

Abstract:

Casing soil is essential for the mushroom formation of Agaricus bisporus, which limits the efficient development of the mushroom industry. This paper reviews the research progress on the mechanisms of ethylene and 1-octen-3-ol on the casing-induced mushroom formation of Agaricus bisporus, including the synthesis and action mechanism of the volatile self-inhibitors for the mushroom formation, and prospects for future research directions.

Key words: Agaricus bisporus, ethylene, 1-octen-3-ol, casing soil, mechanism of the mushroom formation

张朝辉, 闫鹏, 张广, 文瑾瑾, 武忠伟, 王振河, 邱立友. 双孢蘑菇覆土出菇机理研究进展[J]. 园艺学报, 2023, 50(9): 2048-2058.

ZHANG Chaohui, YAN Peng, ZHANG Guang, WEN Jinjin, WU Zhongwei, WANG Zhenhe, QIU Liyou. Research Progress on the Mechanism of the Mushroom Formation in the Button Mushroom（Agaricus bisporus）Induced by Casing Soils[J]. Acta Horticulturae Sinica, 2023, 50(9): 2048-2058.

图/表 4

图1 双孢蘑菇乙烯合成途径黑色箭头表示乙烯合成路径。Met：甲硫氨酸；SAMS：S-腺苷甲硫氨酸合成酶；SAM：S-腺苷甲硫氨酸；MTA：5’-甲硫腺苷；MTR：5’-甲硫基核糖；MTR-1P：5-甲硫核糖1-磷酸；KMBA：2-酮-4-甲基硫代丁酸；ACS：ACC合成酶；ACC：1-氨基环丙烷-1-羧酸；ACO：ACC氧化酶。

Fig. 1 Ethylene synthesis pathway of Agaricus bisporus The black arrow indicates the ethylene synthesis path；Met：Methionine；SAMS：S-adenosine methionine synthase；SAM：S-adenosine methionine；MTA：5’-methylthioadenosyl；MTR：5’-methylthioribose；MTR-1P：5-methylthioribose-1-phosphate；KMBA：α-keto-γ-methylthiobutyric acid；ACS：ACC synthase；ACC：1-amino cyclopropane-1-carboxylic acid；ACO：ACC oxidase.

图2 ACC脱氨酶产生菌解除乙烯抑制双孢蘑菇子实体形成的机理黑色箭头表示乙烯合成路径，红色箭头表示乙烯抑制子实体的分化和形成。Met：甲硫氨酸；SAMS：S-腺苷甲硫氨酸合成酶；SAM：S-腺苷甲硫氨酸；ACS：ACC合成酶；ACC：1-氨基环丙烷-1-羧酸；ACO：ACC氧化酶；Acds：含1-氨基环丙烷-1-羧酸脱氨酶的细菌；FU：菌索；PD：原基；DP：已分化子实体；FB：子实体。

Fig. 2 The mechanism of ACC deaminase-producing bacteria relieving the inhibition of ethylene on the formation of fruiting body of Agaricus bisporus The black arrow indicates the pathway of ethylene synthesis，and the red arrow indicates that ethylene inhibits the differentiation and formation of fruiting bodies. Met：Methionine；SAMS：S-adenosine methionine synthase；SAM：S-adenosylmethionine；ACS：ACC synthase；ACC：1-Amino cyclopropane-1-carboxylic acid；ACO：ACC oxidase；Acds：Microorganisms rich in 1-aminocyclopropane-1-carboxylic acid deaminase；FU：Mycelium knot；PD：Primordium；DP：Differentiated fruiting body；FB：Fruiting body.

表1 双孢蘑菇栽培过程培养料中、覆土中和子实体上微生物群落的多样性

Table 1 Microbial community diversity in the compost，casing soil and on mushrooms during the process of composting and Agaricus bisporus cultivation

生产阶段Crop stage	培养料Compost	覆土层Casing layer	子实体Fruiting body	参考文献Reference
培养料发酵 Compost fermentation	嗜热双歧菌Thermobifida			Carrasco et al.,2019
	嗜热芽孢杆菌属Thermobacillus			Song et al.,2021
	瘤胃梭菌属Ruminiclostridium			隽加香等,2019
	芽胞杆菌属Bacillus			Braat et al.,2022
菌丝生长 Mycelial growth	螯台球菌属Chelatococcus			Carrasco et al.,2020
	栖热菌属Thermus			Carrasco et al.,2020
	未知属Themofibia			Carrasco et al.,2020
覆土 Casing	假单胞菌属Pseudomonas	假单胞菌属Pseudomonas		Rainey,1991a,1991b
	栖热菌属Thermus	黄杆菌属Flavobacterium		Taparia et al.,2021
	嗜热双孢菌属Thermobispora	鞘氨醇杆菌Sphingobium		Carrasco et al.,2019； Braat et al.,2022
	热脱硫杆菌属Thermodesulfobacterium	假黄色单胞菌属Pseudoxanthomonas		McGee et al.,2017
菌丝扭结 Hyphae kink	假单胞菌属Pseudomonas	假单胞菌属Pseudomonas		Chen et al.,2013； Braat et al.,2022
	小梨形菌属Pirellula	黄杆菌属Flavobacterium		Carrasco et al.,2019； Braat et al.,2022
	嗜热双歧菌Thermobifida	蛭弧菌属Bdellovibrio		Carrasco et al.,2019； Braat et al.,2022
出菇 Fruiting	假单胞菌属Pseudomonas	假单胞菌属Pseudomonas	假单胞菌属Pseudomonas	Li et al.,2020； Braat et al.,2022
	小梨形菌属Pirellula	黄杆菌属Flavobacterium	不动杆菌属Acinetobacter	Carrasco et al.,2019； Braat et al.,2022
	栖热菌属Thermus	蛭弧菌属Bdellovibrio	黄杆菌属Flavobacterium	Carrasco et al.,2019； Braat et al.,2022

图3 乙烯影响双孢蘑菇子实体形成的机制紫色箭头表示诱导作用，黑色箭头表示生理过程，绿色箭头表示促进作用，红色箭头表示抑制作用。FU：菌索；DP：已分化子实体；NOX：NADPH氧化酶；Lac：漆酶；MnP：锰过氧化物酶；ROS：活性氧。

Fig. 3 The mechanism of the influence of ethylene on development of Agaricus bisporus The purple arrow represents induction，the black arrow shows the physiological process，the green arrow represents facilitation，and the red arrow represents inhibition. FU：Mycelium knot；DP：Differentiated fruiting body；NOX：NADPH oxidase；Lac：laccase；MnP：Manganese peroxidase；ROS：Reactive oxygen species.

参考文献 75

[1]	Allen C J, Lacey R F, Binder Bickford A B, Beshears C P, Gilmartin C J, Binder B M. 2019. Cyanobacteria respond to low levels of ethylene. Frontiers in Plant Science, 10:950. doi: 10.3389/fpls.2019.00950 pmid: 31417582
[2]	Baars J J P, Scholtmeijer K, Sonnenberg A S M, Peer A V. 2020. Critical factors involved in primordia building in Agaricus bisporus:a review. Molecules, 25 (13):2984. doi: 10.3390/molecules25132984 URL
[3]	Bidon B, Kabbara S, Courdavault V, Glévarec G, Oudin A, Héricourt F, Carpin S, Spíchal L, Binder B M, Cock J M, Papon N. 2020. Cytokinin and ethylene cell signaling pathways from prokaryotes to eukaryotes. Cells, 9 (11):2526. doi: 10.3390/cells9112526 URL
[4]	Bonnen A M, Anton L H, Orth A B. 1994. Lignin-degrading enzymes of the commercial button mushroom, Agaricus bisporus. Applied and Environmental Microbiology, 60 (3):960-965.
[5]	Braat N, Koster M C, Wösten H A B. 2022. Beneficial interactions between bacteria and edible mushrooms. Fungal Biology Reviews, 39:60-72. doi: 10.1016/j.fbr.2021.12.001 URL
[6]	Buechner R, Voros M, Allaga H, Varga A, Bartal A, Szekeres A, Varga S, Bajzat J, Bakos-Barczi N, Misz A, Csutoras C, Hatvani L, Vagvolgyi C, Kredics L. 2022. Selection and characterization of a bacillus strain for potential application in industrial production of white button mushroom(Agaricus bisporus). Agronomy-basel, 12 (2):467.
[7]	Byrne A R, Tušek Žnidarič M. 1990. Studies of the uptake and binding of trace metals in fungi. Part I:accumulation and characterization of mercury and silver in the cultivated mushroom, Agaricus bisporus. Applied Organometallic Chemistry, 4 (1):43-48.
[8]	Cano-Dominguez N, Alvarez-Delfin K, Hansberg W, Aguirre J. 2008. NADPH oxidases NOX-1 and NOX-2 require the regulatory subunit nor-1 to control cell differentiation and growth in Neurospora crassa. Eukaryotic Cell, 7 (8):1352-1361. doi: 10.1128/EC.00137-08 pmid: 18567788
[9]	Carrasco J, Garcia-Delgado C, Lavega R, Tello M L, de Toro M, Barba-Vicente V, Rodriguez-Cruz M S, Sanchez-Martin M J, Perez M, Preston G M. 2020. Holistic assessment of the microbiome dynamics in the substrates used for commercial champignon(Agaricus bisporus)cultivation. Microbial Biotechnology, 13 (6):1933-1947 doi: 10.1111/mbt2.v13.6 URL
[10]	Carrasco J, Tello M L, De-Toro M, Tkacz A, Poole A, Pérez-Clavijo M, Preston M. 2019. Casing microbiome dynamics during button mushroom cultivation: implications for dry and wet bubble diseases. Microbiology, 165:611-624. doi: 10.1099/mic.0.000792
[11]	Chagué V. 2010. Ethylene Production by Fungi:Biological questions and future developments towards a sustainable polymers industry. Handbook of hydrocarbon and lipid microbiology. Handbook of hydrocarbon and lipid microbiology.Berlin:Springer-Verlag Berlin Heidelberg.
[12]	Chen J Q, Chen T C, Qiu M M, Li L, Zhong Q G, Wei Q, Deng Y J, Xie B G, Jiang Y J, Chen B Z. 2021. Identification of ACC synthetase genes in Volvariella volvacea and analysis of their response to ethephon and 1-methylcyclopropene treatments. Scientia Horticulturae, 278:109848. doi: 10.1016/j.scienta.2020.109848 URL
[13]	Chen S C, Qiu C W, Huang T, Zhou W W, Qi Y C, Gao Y Q, Shen J W, Qiu L Y. 2013. Effect of 1-aminocyclopropane-1-carboxylic acid deaminase producing bacteria on the hyphal growth and primordium initiation of Agaricus bisporus. Fungal Ecology, 6 (1):110-118. doi: 10.1016/j.funeco.2012.08.003 URL
[14]	Cheng Yan, Wang Jing-mao, Zhang Yan, Yang Qian-long, Qiu Li-you. 2015. Application of 1-aminocyclopropane-1-carboxylic acid deaminase producing bacterium for increasing the yield of the button mushroom. Henan Science, 33 (10):1750-1755. (in Chinese)
	程雁, 王景冒, 张岩, 杨潜龙, 邱立友. 2015. 应用1-氨基环丙烷-1-羧酸脱氨酶产生菌提高双孢蘑菇产量. 河南科学, 33 (10):1750-1755.
[15]	Chitarra G S, Abee T, Rombouts F M, Posthumus M A, Dijksterhuis J. 2004. Germination of Penicillium paneum conidia is regulated by 1-octen-3-ol,a volatile self-inhibitor. Applied Biochemistry and Microbiology, 70 (5):2823-2829.
[16]	Dubey A K, Suresh C, Umesh Kumar S, Karanth N G. 1998. An enzyme-linked immunosorbent assay for the estimation of fungal biomass during solid-state fermentation. Applied Microbiology and Biotechnology, 50 (3):299-302. pmid: 9802214
[17]	Eastwood D C, Herman B, Noble R, Dobrovin-Pennington A, Sreenivasaprasad S, Burton K S. 2013. Environmental regulation of reproductive phase change in Agaricus bisporus by 1-octen-3-ol,temperature and CO₂. Fungal Genetics and Biology, 55:54-66. doi: 10.1016/j.fgb.2013.01.001 pmid: 23354075
[18]	Elisashvili V, Kachlishvili E, Penninckx M. 2008. Lignocellulolytic enzymes profile during growth and fruiting of Pleurotus ostreatus on wheat straw and tree leaves. Acta Microbiologica et Immunologica Hungarica, 55 (2):157-168. doi: 10.1556/AMicr.55.2008.2.7 pmid: 18595320
[19]	Fukuda H, Ogawa T, Tanase S. 1993. Ethylene production by micro-organisms. Academic Press.:275-306.
[20]	Gessler N N, Filippovich S Y, Bachurina G P, Kharchenko E A, Groza N V, Belozerskaya T A. 2017. Oxylipins and oxylipin synthesis pathways in fungi. Applied Biochemistry and Microbiology, 53 (6):628-639. doi: 10.1134/S0003683817060060 URL
[21]	Guo H, Liu A R, Wang Y R, Wang T, Zhang W, Zhu P K, Xu L. 2020. Measuring light-induced fungal ethylene production enables non-destructive diagnosis of disease occurrence in harvested fruits. Food Chemistry, 310:125827. doi: 10.1016/j.foodchem.2019.125827 URL
[22]	Holighaus G, Rohlfs M. 2019. Volatile and non-volatile fungal oxylipins in fungus-invertebrate interactions. Fungal Ecology, 38:28-36. doi: 10.1016/j.funeco.2018.09.005
[23]	Juan Jia-xiang, Xiao Ting-ting, Wang Qian, Song Xiao-xia, Shen Xin-fen, Gao Fei, Chen Ming-jie, Huang Jian-chun. 2019. Microbial population during composting of Agaricus bisporus culture substrate and functional prediction thereof. Acta Edulis Fungi, 26 (4):50-56,159-160. (in Chinese)
	隽加香, 肖婷婷, 王倩, 宋晓霞, 沈新芬, 高飞, 陈明杰, 黄建春. 2019. 双孢蘑菇发酵培养料细菌菌群结构及其功能预测. 食用菌学报, 26 (4):50-56,159-160.
[24]	Ke De-sen, Wang Zheng-xun. 2009. The role of reactive oxygen species in the ethylene biosynthesis in plants. Journal of Wuhan Botancial Research, 27 (3):327-331. (in Chinese)
	柯德森, 王正询. 2009. 植物乙烯生物合成过程中活性氧的作用. 武汉植物学研究, 27 (3):327-331.
[25]	Kim K, Kim A. 2005. Optimization for effective bioproduction of natural (-)-1-Octen-3-ol by lipoxygenase and hydroperoxide lyase from Agaricus bisporus. Journal of the Korean Society of Food Science and Nutrition, 34:899-903. doi: 10.3746/jkfn.2005.34.6.899 URL
[26]	Lee N Y, Choi D H, Kim M G, Jeong M J, Kwon H J, Kim D H, Kim Y G, di Luccio E, Arioka M, Yoon H J, Kim J G. 2020. Biosynthesis of (R)-(-)-1-Octen-3-ol in recombinant saccharomyces cerevisiae with lipoxygenase-1 and hydroperoxide lyase genes from Tricholoma matsutake. Journal of Microbiology and Biotechnology, 30 (2):296-305. doi: 10.4014/jmb.2001.01049 URL
[27]	Li J, Wei Q, Huang L X, Fang T, Chen B Z, Jiang Y J. 2020. Mathematical modeling Pseudomonas spp. growth and microflora composition variation in Agaricus bisporus fruiting bodies during chilled storage. Postharvest Biology and Technology, 163:111144. doi: 10.1016/j.postharvbio.2020.111144 URL
[28]	Li T, Zhang J, Shen C H, Li H R, Qiu L Y. 2019. 1-Aminocyclopropane-1-carboxylate:a novel and strong chemoattractant for the plant beneficial rhizobacterium Pseudomonas putida UW4. Molecular Plant-microbe Interactions, 32 (6):750-759. doi: 10.1094/MPMI-11-18-0317-R URL
[29]	Li W H, Li Q Y, Lyu M, Wang Z J, Song Z H, Zhong S W, Gu H Y, Dong J, Dresselhaus T, Zhong S, Qu L J. 2022. Lack of ethylene does not affect reproductive success and synergid cell death in Arabidopsis. Molecular Plant, 15 (2):354-362. doi: 10.1016/j.molp.2021.11.001 URL
[30]	Li Ya-nan, Zhang Yan, Shang Di, Gao Yu-qian, Liu Qin, Cui Xiao, Kong Wei-li, Qiu Li-you. 2021. Ammonia tolerance of different Pleurotus ostreatus strains. Acta Edulis Fungi, 28 (4):57-63. (in Chinese)
	李亚楠, 张艳, 尚迪, 高玉千, 刘芹, 崔筱, 孔维丽, 邱立友. 2021. 不同糙皮侧耳菌株氨耐受性分析. 食用菌学报, 28 (4):57-63.
[31]	Long P E, Jacobs L. 1974. Aseptic fruiting of the cultivated. Transactions of the British Mycological Society, 63 (1):99-107. doi: 10.1016/S0007-1536(74)80140-3 URL
[32]	Macedo G E, Brum V P D, Rodrigues N R, Gomes K K, Martins I K, Franco J L, Posser T. 2020. Fungal compound 1-octen-3-ol induces mitochondrial morphological alterations and respiration dysfunctions in Drosophila melanogaster. Ecotoxicology and Environmental Safety, 206:111232. doi: 10.1016/j.ecoenv.2020.111232 URL
[33]	Mau J, Beelman R B, Ziegler G R. 1992. Effect of 10-oxo-trans-8-decenoic acid on growth of Agaricus bisporus. Phytochemistry, 31 (12):4059-4064. doi: 10.1016/0031-9422(92)80414-A URL
[34]	McGee C F, Byrne H, Irvine A, Wilson J. 2017. Diversity and dynamics of the DNA and cDNA-derived bacterial compost communities throughout the Agaricus bisporus mushroom cropping process. Annals of Microbiology, 67 (11):751-761. doi: 10.1007/s13213-017-1303-1 URL
[35]	Mehmood A, Liu G R, Wang X, Meng G N, Wang C T, Liu Y. 2019. Fungal quorum-sensing molecules and inhibitors with potential antifungal activity:a review. Molecules, 24 (10):1950. doi: 10.3390/molecules24101950 URL
[36]	Mu D S, Li C Y, Zhang X C, Li X B, Shi L, Ren A, Zhao M W. 2014. Functions of the nicotinamide adenine dinucleotide phosphate oxidase family in Ganoderma lucidum:an essential role in ganoderic acid biosynthesis regulation,hyphal branching,fruiting body development,and oxidative-stress resistance. Environmental Microbiology, 16 (6):1709-1728. doi: 10.1111/emi.2014.16.issue-6 URL
[37]	Muszyńska B, Kała K, Rojowski J, Grzywacz A, Opoka W. 2017. Composition and biological properties of Agaricus bisporus fruiting bodies- a review. Polish Journal of Food and Nutrition Sciences, 67 (3):173-181. doi: 10.1515/pjfns-2016-0032 URL
[38]	Noble R, Dobrovin-Pennington A, Hobbs P J, Pederby J, Rodger A. 2009. Volatile C 8 compounds and pseudomonads influence primordium formation of Agaricus bisporus. Mycologia, 101 (5):583-591. doi: 10.3852/07-194 URL
[39]	Noble R, Fermor T R, Lincoln S, Dobrovin-Pennington A, Evered C, Mead A, Li R. 2003. Primordia initiation of mushroom(Agaricus bisporus) strains on axenic casing materials. Mycologia, 95 (4):620. pmid: 21148971
[40]	North J A, Narrowe A B, Xiong W, Byerly K M, Zhao G, Young S J, Murali S, Wildenthal J A, Cannon W R, Wrighton K C, Hettich R L, Tabita F R. 2020. A nitrogenase-like enzyme system catalyzes methionine,ethylene,and methane biogenesis. Science, 369 (6507):1094-1098. doi: 10.1126/science.abb6310 URL
[41]	Ohga S. 1998. Carbon dioxide and ethylene levels during incubation and fruiting stages on sawdust-based culture of Lentinula edodes. Kyushu University Forests, 79:13-20.
[42]	Ohga S, Royse D J. 2001. Transcriptional regulation of laccase and cellulase genes during growth and fruiting of Lentinula edodes on supplemented sawdust. FEMS Microbiology Letters, 201 (1):111-115. pmid: 11445176
[43]	Park J Y, Agnihotri V P. 1969. Bacterial metabolites trigger sporophore formation in Agaricus bisporus. Nature, 222 (5197):984.
[44]	Polat E, Erler F, Demir H, Cetin H. 2008. The effect of vegetable materials on the yield and productivity of Agaricus bisporus. Interciencia Journal, 33 (10):776-780.
[45]	Rainey P B. 1991a. Effect of Pseudomonas putida on hyphal growth of Agaricus bisporus. Mycological Research, 95 (6):699-704. doi: 10.1016/S0953-7562(09)80817-4 URL
[46]	Rainey P B. 1991b. Phenotypic variation of Pseudomonas putida and P. tolaasii affects attachment to Agaricus bisporus mycelium. Journal of General General Microbiology Microbiology, 137 (12):2769-2779.
[47]	Ralph H K J. 1995. Cobalt chloride and ethylene affect fruiting of Agaricus bisporus. Mycologia, 3 (87):366-369.
[48]	Ramoni R, Vincent F, Grolli S, Conti V, Malosse C, Boyer F, Meillour P N, Spinelli S, Cambillau C, Tegoni M. 2001. The insect attractant 1-Octen-3-ol is the natural ligand of bovine odorant-binding protein. Journal of Biological Chemistry, 276 (10):7150-7155. doi: 10.1074/jbc.M010368200 pmid: 11114310
[49]	Riyazuddin R, Verma R, Singh K, Nisha N, Keisham M, Bhati K K, Kim S T, Gupta R. 2020. Ethylene:a master regulator of salinity stress tolerance in plants. Biomolecules, 10 (6):959. doi: 10.3390/biom10060959 URL
[50]	Scott B, Eaton C J. 2008. Role of reactive oxygen species in fungal cellular differentiations. Current Opinion Microbiology, 11 (6):488-493. doi: 10.1016/j.mib.2008.10.008 URL
[51]	Singh D, Son S Y, Lee C H. 2020. Critical thresholds of 1-Octen-3-ol shape inter-species Aspergillus interactions modulating the growth and secondary metabolism. Scientific Reports, 10 (1):1-14. doi: 10.1038/s41598-019-56847-4
[52]	Song T T, Shen Y Y, Jin Q L, Feng W L, Fan L J, Cao G T, Cai W M. 2021. Bacterial community diversity,lignocellulose components,and histological changes in composting using agricultural straws for Agaricus bisporus production. Peerj, 9:e10452.
[53]	Stark W, Forss D A. 1964. A compound responsible for mushroom flavour in dairy products. Journal of Dairy Research, 31 (3):253-259. doi: 10.1017/S0022029900018185 URL
[54]	Stoller B. 1952. Studies on the function of the casing soil for mushroom beds. Mushroom Grow Assoc Bull, 4 (34):289-297.
[55]	Sun L B, Xin G, Hou Z S, Zhao X M, Xu H R, Bao X J, Xia R R, Li Y T, Li L. 2021. Biosynthetic mechanism of key volatile biomarkers of harvested Lentinula edodes triggered by spore release. Journal of Agricultural and Food Chemistry, 69 (32):9350-9361. doi: 10.1021/acs.jafc.1c02410 URL
[56]	Taparia T, Hendrix E, Hendriks M, Nijhuis E, Boer D W, Wolf J V D. 2021. Casing soil microbiome mediates suppression of bacterial blotch of mushrooms during consecutive cultivation cycles. Soil Biology and Biochemistry, 155:108161. doi: 10.1016/j.soilbio.2021.108161 URL
[57]	Tsitsigiannis D I, Kowieski T M, Zarnowski R, Keller N P. 2004. Endogenous lipogenic regulators of spore balance in Aspergillus nidulans. Eukaryotic Cell, 3 (6):1398-1411. pmid: 15590815
[58]	Turner E M, Wright M, Ward T, Osborne D J, Self R. 1975. Production of ethylene and other volatiles and changes in cellulase and laccase activities during the life cycle of the cultivated mushroom, Agaricus bisporus. Journal of General Microbiology, 91 (1):167.
[59]	Wang K L C, Li H, Ecker J R. 2002. Ethylene biosynthesis and signaling networks. The Plant Cell, 14 (suppl 1):S131-S151. doi: 10.1105/tpc.001768 URL
[60]	Ward T, Turner E M, Osborne D J. 1978. Evidence for the production of ethylene by the mycelium of Agaricus bisporus and its relationship to sporocarp development. Microbiology, 104:23-30.
[61]	Wei Peng-fei, Chen Jian-fang, Yang Qian-long, Zheng Yun-feng, Qiu Li-you, Gao Yu-qian. 2019. Function of the casing materials for mushroom beds. Northen Horticulture,(18):133-139. (in Chinese)
	魏鹏飞, 陈建芳, 杨潜龙, 郑云峰, 邱立友, 高玉千. 2019. 双孢蘑菇高产栽培覆土新技术. 北方园艺 (18):133-139.
[62]	Wood D A, Blight M. 1982. Sporophore initiation in axenic culture. Report of the Glasshouse Crops Research Institute for 1981. 140.
[63]	Wood D A, Hammond J B. 1977. Ethylene production by axenic fruiting cultures of Agaricus bisporus. Applied and Environmental Microbiology, 34 (2):228-229. doi: 10.1128/aem.34.2.228-229.1977 pmid: 562129
[64]	Wurzenberger M, Grosch W. 1982. The enzymic oxidative breakdown of linoleic acid in mushrooms (Psalliota bispora). Zeitschrift für Lebensmittel-Untersuchung und Forschung, 175 (3):186-190. doi: 10.1007/BF01139769 URL
[65]	Wurzenberger M, Grosch W. 1983. Bestimmung von 1-Oeten-3-ol in Pilzen und Pilzprodukten. Zeitschrift für Lebensmittel-Untersuchung und Forschung, 176 (1):16-19. doi: 10.1007/BF01089340 URL
[66]	Xiong C, Li Q, Li S H, Chen C, Chen Z Q, Huang W L. 2017. In vitro antimicrobial activities and mechanism of 1-Octen-3-ol against food-related bacteria and pathogenic fungi. Journal of Oleo Science:1041-1049.
[67]	Yang C, Lu X, Ma B, Chen S Y, Zhang J S. 2015. Ethylene signaling in rice and Arabidopsis:conserved and diverged aspects. Molecular Plant, 8 (4):495-505. doi: 10.1016/j.molp.2015.01.003 URL
[68]	Zawirska-Wojtasiak R. 2004. Optical purity of (R)-(-)-1-octen-3-ol in the aroma of various species of edible mushrooms. Food Chemistry, 86 (1):113-118. doi: 10.1016/j.foodchem.2003.08.016 URL
[69]	Zhang C H, Huang T, Shen C H, Wang X T, Qi Y C, Shen J W, Song A D, Qiu L Y, Ai Y C. 2016. Downregulation of ethylene production increases mycelial growth and primordia formation in the button culinary-medicinal mushroom, Agaricus bisporus(Agaricomycetes). International Journal of Medicinal Mushrooms, 18 (12):1131-1140.
[70]	Zhang C H, Zhang G, Wen Y M, Li T, Gao Y Q, Meng F M, Qiu L Y, Ai Y C. 2019. Pseudomonas sp. UW4 acdS gene promotes primordium initiation and fruiting body development of Agaricus bisporus. World Journal of Microbiology and Biotechnology, 35 (11):163. doi: 10.1007/s11274-019-2741-7
[71]	Zhang Chao-hui, Zhang Guang, Wen Ya-mei, Yang Yan-qing, Liu Ya-fei, Zhang Hui, Wang Zhen-he, Qiu Li-you. 2019. The role of 1-aminocyclopropane-1-carboxylate in the chemotactic response of Pseudomonas putida to Agaricus bisporus mycelia. Acta Agriculturae Boreali-Sinica, 34 (6):209-218. (in Chinese)
	张朝辉, 张广, 闻亚美, 杨艳青, 刘亚非, 张会, 王振河, 邱立友. 2019. 1-氨基环丙烷-1-羧酸在恶臭假单胞菌趋化双孢蘑菇菌丝中的作用. 华北农学报, 34 (6):209-218. doi: 10.7668/hbnxb.20190496
[72]	Zhang Da-fei, Qi Yuan-cheng, Gao Yu-qian, Shen Jin-wen, Qiu Li-you. 2010. A primary analysis on the mechanism of casing soil triggering the sporophore formation of Agaricus bisporus. Edible Fungi, 32 (1):9-11. (in Chinese)
	张大飞, 戚元成, 高玉千, 申进文, 邱立友. 2010. 双孢蘑菇覆土出菇机理初步探讨. 食用菌, 32 (1):9-11.
[73]	Zhang Yan, Zhang Chao-hui, Liu Dong-ren, Wang Xiao-fei, Wang Jing-mao, Qiu Li-you. 2015. Liposome- mediated Fruit body gill tissue transformation of Agaricus bisporus and function exploration of 1-aminocyclopropane-1-carboxylic acid oxidase gene(ACO). Journal of Agricultural Biotechnology, 23 (8):1112-1120. (in Chinese)
	张岩, 张朝辉, 刘冬忍, 王小飞, 王景冒, 邱立友. 2015. 脂质体介导遗传转化双孢蘑菇菌褶及1-氨基环丙烷-1-羧酸氧化酶基因(ACO)功能初探. 农业生物技术学报, 23 (8):1112-1120.
[74]	Zhao K, Zhu X, Ma H, Ji J, Jin X, Sun J. 2021. Design and experiment of the environment control system for the industrialized production of Agaricus bisporus. International Journal of Agricultural and Biological Engineering, 14 (1):97-107.
[75]	Zuo Y T, Yang Q L, Li Y F, Zhang J, Qi Y C, Gao Y Q, Song A D, Shen J W, Qiu L Y. 2016. Functions and bioinformatics analysis of the NADPH oxidase NoxA in basidiomycete Pleurotus ostreatus:ROS production and fruiting body formation. Academia Journal of Agricultural Research, 4:218-229.