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Ultrastructural Studies on the Cultivation Processes and Growth and Development of the Cultivated Mushroom Agaricus Bisporus

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Ultrastructural Studies on the Cultivation Processes and Growth and Development of the Cultivated Mushroom Agaricus Bisporus Food Structure Volume 4 Number 1 Article 17 1985 Ultrastructural Studies on the Cultivation Processes and Growth and Development of the Cultivated Mushroom Agaricus Bisporus D. A. Wood G. D. Craig P. T. Atkey R. J. Newsam K. Gull Follow this and additional works at: https://digitalcommons.usu.edu/foodmicrostructure Part of the Food Science Commons Recommended Citation Wood, D. A.; Craig, G. D.; Atkey, P. T.; Newsam, R. J.; and Gull, K. (1985) "Ultrastructural Studies on the Cultivation Processes and Growth and Development of the Cultivated Mushroom Agaricus Bisporus," Food Structure: Vol. 4 : No. 1 , Article 17. Available at: https://digitalcommons.usu.edu/foodmicrostructure/vol4/iss1/17 This Article is brought to you for free and open access by the Western Dairy Center at DigitalCommons@USU. It has been accepted for inclusion in Food Structure by an authorized administrator of DigitalCommons@USU. For more information, please contact [email protected]. FOOD MICROSTRUCfURE, Vol. 4 (1985). pp. 143-164 0730-5419/85$1 . 00+.05 SEM Inc .. AMF O'Hare (Chicago) , IL 60666-0507 U.S.A. ULTRASTRUCTURAL STUDIES ON THE CULTIVATION PROCESSES AND GROWTH AND DEVELOPMENI' OF THE CULTIVATED MUSHROOM AGARICUS BISPORUS 1 2 1 3 3 D.A. Wood *, G.D. Craig , P. T. Atkey , R.J. Newsarn and K. G.Jll 1 Dept. of Plant Pathology & Microbiology , Glasshouse Crops Research Institute, Littlehampton, 2 West Sussex, BN17 6LP, UK 3 Dept. of Life Sciences, Nottingham Polytechnic, Nottingham, UK Biological Laboratory, University of Kent, canterbury, Kent, UK Abstract Introduction Scanning electron microscopy (SEM) , The production of the cultivated mushroom is transmission electron microscopy and light carried out on a large scale in North America, microscopy have been used t o study various Europe, Australasia and parts of Asia . World processes in the cultivation of the edible production now exceeds 1.0 million t onnes per mushroom Agaricus bisporus . Mushrooms are annum. Production is carried out by growing the cultured on composted wheat straw. The microbial mycelium of the fungus in composted substrates, degradation processes during composting have been such as manure-straw mixtures , a nd then harvest­ visualised by TEM and SEM and correlated with ing the mushroom fruit bodies prior t o marketing microbiological studies. Various modes of attack (Wood, 1984, Wood et al., 1984). In the United by the microorganisms on the plant cell walls can Kingdom the consumer preference is f or fresh be seen. Most rapid degradation occurred on the mushrooms but in North America, France and Asia cuticle and phloem and spread to other cell considerable quantitites of mushrooms are canned types. Microbial attack was found to be non­ before marketing . uniform between tissue types and individual Many microbiological studies have been cells. The mycelium of Agaricus bisporus carried out on the composting process used for colonised the compost straw surface and the lumen the growth substrote on which mushrooms are of straw cells, and also degraded microbial cells cultivated (Fermor and Wood, 1979) . A large and therein. taxonomically diverse microbial flora has been The cellular organisation and ultrastructure shown to be involved in composting . These of the rhizomorphs and of the developing stipe microbes include bacteria, actinomycetes and and gill tissue were examined. The development fungi and both mesophilic and thermophilic and branching patterns of the hyphae generating microbial g roups a re found. The soluble the sub-hymenium and hymenium l ayer s of the gill materials a nd certain of the insoluble materials tissue were examined . Light and electron of the substrate are utilised during composting microscope autoradiography were used t o locate (Fermor and Wood, 1979 , Fermor a nd Wood, 1981). the site of synthesis of cell wall chitin and to After composting , the growth substrate demonstrate evidence for cell division in the (compost) is inoculated with mushroom mycelium upper stipe region. and the compos t is subsequently extensively colonised by the mycelium . Mushrooms arise as small knots or aggregates of hyphal cells and differentiate into the 3 tissue types of the Initial paper received February 18, 1985. mature structure, stipe, cap and gills. Final manuscript received May 21, 1985. We investigated the ultrastructure of the Direct inquiries to D.A. Wood. composting process and colonisation stages of Telephone number: (0903) 716123. mushroom production t o determine the organisation and mode of attack of the microorganisms in the compost and t o study the colonisation by the hyphae of the mushroom into the substrate. The ~ey_~-r~: ITRlShrcx:rn, Agar~~, scanning electron development of the mushroom from the earliest microscopy, transmission electron microscopy, stages to mature fruit bodies was examined by ultrastructure, ccrrp::Jst, growth, developrent, transmission e lec tron microscopy (TEM) and autoradiography, cell division. scanning electron microscopy (SEM). Auto­ radiographic techniques were used to examine cell division and chitin synthesis in stipe tissue of the mushrooms. 143 D. A. Wood , G. D. Craig , P. T. Atkey, et al . Pha se 1 - Composting of straw/manures ~ 3 Phase 2 - Composting, high temperature stage + In oculation o f compost Colonisation Application of casing material ~ First fruit bodies nC ~ Primordium Set of fruiting cycles ~ End of crop G Fig . 1 A simplified schematic f l ow chart of the main stages in mushroom cultivation Fig. 4 Day 0 . SEM of raw straw surface. Note B silica l:x:xlies (arrows) . Bar 50 11m . ~h_ Hyphal Fig. 5 Day 0. Sil1 of straw surface fran stable ITI3Ilure. tJote diverse microbial population. Bar 50 11m . -- knot Fig. 6 Day 0 . SEM of cut end of straw internode / from stable ITI3Ilure showing microbial concentration in phloan. Bar 50 11rn. Fig. 7 Day 7. 'IEM of variety of microorganisms Initiation within a straw cell. Bar 2 . 5 1111' . Fig. 8 Day 7 . Til1 of straw showing absence of microbial colonisation. Bar 5 11m . Fig. 9 Day 7 . SEM of col ony of coccoid bacterial Fruiting of cells on the straw surface. Bar 5 l.llll. Fig. 10 Day 7 . SEM of microorganisms on straw ~garicus bisr:2orus cuticle. Note erosion aroW1d silica l:x:xly (arrow). Bar 25 l.llll. Morphology Fig. ll Day 14 . Til'l of bacteria within straw cells. Note erosion (W , cell wall) and glycocalyx Fig . 3 Diagram of morphogenesis of the mushroom rraterial anchoring bacteria (g) . Bar 1 11m . Agaricus bisporus Fi g . 2 Diagram of growth a nd cropping of the edible mushroom Agaricus bisporus on composted substrates Development of the Cultivated Mushroom Open cup (section) pileus (cap) 2 144 Ultrastructural Studies on Mushroom Cultivation 145 D.A. Wood, G.D. Craig, P.T. Atkey, et al. Materials and Methods T20 scanning microscope at 20 kV accelerating voltage. Samples from mushroom cultivation processes A schemat1c flow chart of the major steps in Samples from mushroom tissue mushroom cultivation is shown in Figure 1. A A diagram of the morphogenesis of Agaricus more detailed description of the process and the bisporus is shown in Figure 3. Samples for environmental and cultural parameters controlling microscopy were taken from stages c, E, F, G. it and their purpose can be found in Wood (1984). A summary of the growing procedure for mushrooms Light microscopy is shown in Fig. 2. Samples of straw internode -----Small pieces of stipe tissue were removed at material were taken from the bulk of the compost stages F and G, and fixed in 2.5% glutaraldehyde at each of the following stages of the in sodium cacodylate pH 7.4. The samples were cultivation process. Raw straw + stable manure post fixed in 1% osmium tetroxide in veronal­ at 0 d , first phase of composting at 7 d, acetate buffer pH 7.2 and then dehydrated in an pasteurization at 14 d; mycelial inoculation at ethanol series and embedded in Spurr's resin 21 d, application of casing material at 35 d, (Spurr, 1969). The tissue was embedded so first appearance of fruit bodies at 56 d and end longitudinal sections could be cut to reveal the of fruiting at 112 d. orientation of the hyphae. Sections (0.5 pm) Transmission electron microscopy (TEM) of straw were cut with glass knives with a Reichert OMU3 samples ultratome, stained with 1% toludine blue and Pieces of tissue ca 0.5 mm x 2 mm were cut examined in a Zeiss Universal light microscope. from the internode region of the straw. These were fixed for 2 h in Karnovsky's fixative TEM of mushroom tissue (Karnovsky, 1965) using half strength para­ ----samples of the material embedded for light formaldehyde. Samples were fixed in embedding microscopy were sectioned as described above and tubes as described by Pegler and Atkey (1978). then post-stained with uranyl acetate and lead The tissues were thoroughly rinsed in 0.2 M citrate. The sections were examined in an AEI sodium cacodylate buffer pH 7.2; and post-fixed 801 transmission microscope at 60 kV accelerating for 2 h in 2% osmium tetroxide. The tissue was voltage. then washed and dehydrated through a graded ethanol series and embedded in Spurr's resin SEM of mushroom tissue (Spurr, 1969) after infiltration at 24 h for 3 ---~ngitudinal and trans~erse sections of increasing concentrations and a final 48 h in stipe and gill tissue (5 mm ) were removed and 100% resin. Fixation and dehydration was carried fixed for 18 h in 2.5% glutaraldehyde in sodium out at room temperature. Sections were cut with cacodylate pH 7.4. The specimens were dehydrated an LKB Ultratome III using glass knives, and then through a graded acetone series each at 30 stained with aqueous uranyl acetate for 30 minutes and finally into 100% acetone . The minutes and post-stained with Reynolds lead specimens were then critical point dried with citrate solution (Reynolds , 1963) and examined in liquid carbon dioxide with a Polaron model E300 a JSM 100 S transmission microscope at 80 kV drier.
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