Spent substrate from mushroom cultivation: exploitation potential toward various applications and value-added products

Figures & data

Figure 1. Factors affecting cultivation parameters and the use of SMS in new mushroom crops.

Table 1. Reuse of spent mushroom substrate (SMS) for the cultivation of various mushroom species as reported in pertinent publications: origin and composition of SMS, mushroom to be cultivated, new substrate formulation and supplements, biological efficiency (BE) reported for the crop obtained and main comments on the results of the respective study. Abbreviation used: NR, not reported.

Origin of SMS	Composition of SMS	Mushroom to be cultivated	New substrate formulation	Supplements to new substrate	Biological efficiency	Comments	References
Pleurotus ostreatus	NR	P. ostreatus	SMS (plain, 3:1, 1:1 and 1:3) with rubber sawdust	CaCO3 (1.5% w/w), wheat bran (5% w/w)	53–56%	Similar BE values with the control (rubber sawdust; BE: 53%)	[29]
Pholiota nameko, Hypsizigus marmoreus, Hericium erinaceus	Oak and beech sawdust (1:1), wheat bran (20%), corn flour (5%) and gypsum (1%)		SMS and wheat straw (1:9, 2:8 and 3:7)	Wheat bran (3%), gypsum (1%)	66–73%	Reduction in BE was noted when SMS ratios increased	[30]
		P. ostreatus
P. ostreatus	Wheat straw, wheat bran, soybean flour and CaCO3	P. ostreatus	SMS	Wheat bran, soybean flour, and CaCO3 (C/N: 20)	185%	High (the highest reported) BE values for both species examined by using SMS as cultivation substrate	[26]
		Pleurotus pulmonarius			208%
Cyclocybe cylindracea, Auricularia polytricha, H. erinaceus, Lyophyllum shimeji, Pleurotus citrinopileatus, Pleurotus cystidiosus, Pleurotus eryngii, P. ostreatus, Pleurotus sajor-caju	Sawdust	A. polytricha	SMS from various species	Rice bran (9.5%), CaCO3 (.5%)	41–59%	All mixtures exhibited higher BE values than the control (BE: 36%)	[28]
Flammulina velutipes	NR	Cordyceps militaris	SMS:wheat bran: rice bran (8:1:1)	Glucose (20 g L^−1) and peptone (5 g L^−1)	35%	Lower yields and cordycepin concentration in SMS-based substrate than in conventional substrate	[31]
P. eryngii	Sugarcane bagasse (50%), cottonseed hulls (20%), wheat bran (20%), cornmeal (5%), soymeal (3%), lime (1%), gypsum (1%)	C. cylindracea	SMS	Sucrose (1%), lime (1%), wheat bran (0, 10, 20%), and Tenebrio molitor feces (0, 10, 20%)	40–63%	Increasing the rate of supplements positively affected the ΒΕ	[32]
P. pulmonarius	Rubber sawdust supplemented with 20% palm oil sludge	C. cylindracea	SMS alone or mixed with rubber sawdust (3:1, 1:1 and 1:3)	Rice bran (5%), Ca(OH)2 (2%), Mg(SO4) (.2%)	25–41%	BE increased by increasing SMS content	[33]
Agaricus bisporus	NR	P. ostreatus	P. ostreatus SMS, and A. bisporus SMS (various ratios)	CaSO4 (5%), gypsum (5%)	NR	P. ostreatus and A. bisporus SMS (3:2 and 1:1) had similar protein and lower ash content than commercial substrate; SMS affected the produced mushrooms’ quality	[34]
P. ostreatus	NR
P. ostreatus	NR	P. ostreatus	SMS or SMS:wheat straw (1:1)	Wheat bran (50 or 100 g kg^−1), Calprozime (20 g kg^−1), gypsum (5%), CaCO3	3–62%	Reduction in BE by increasing wheat bran supplementation; highest BE values were obtained using Calprozime	[35]
H. marmoreus	NR	P. ostreatus	Cottonseed hulls:SMS:wheat bran (various ratios)	Wheat bran (0–18%), lime (1%), gypsum (1%)	36–61%	Reduction in BE when SMS content was increased	[36]
P. ostreatus	NR	P. ostreatus	SMS:sawdust (7:3, 6:4, 5:5, 4:6 and 2:8)	Wheat bran (20%)	78–105%	Reduction in BE when SMS ratio is increased	[37]
		Pleurotus florida			67–89%
NR		P. eryngii	SMS:farmyard manure (1:1) and SMS as casing material (3 cm thick)		49–107%	Enhanced yield on SMS compared to other casing materials and non-casing substrates	[38]
P. eryngii	Sawdust (50%), cottonseed hulls (25%), wheat bran (25%), gypsum and CaCO3	Volvariella volvacea	Composted or non-composted SMS	Limestone (5%)	10–17%	Composted SMS performed better (higher BE) than non-composted one	[39]
F. velutipes	NR	A. bisporus	SMS	Gypsum (2.83%), CaH4P2O8 (2.83%)	29%	Similar BE values with the control (rice straw), but SMS results in shorter production cycles	[40]
P. ostreatus	NR	Agaricus blazei	Sunflower seed hulls:SMS (milled or unmilled)		13.1 – 22.4%	Milled SMS reduced BE	[41]
A. bisporus	Wheat straw and poultry manure-based commercial compost	A. bisporus casing	Sphagnum peat:SMS (4:1, 3:2. 2:3, 1:4) and plain SMS	CaCO3 (100 g L^−1)	53–110%	Lower BE (than the control) when SMS was incorporated in the casing layer	[42]
P. ostreatus	Banana leaf straw	A. blazei	SMS with burned rice husk or subsoil as casing material	CaCO3, urea, rice bran (1, 10 or 20%), (NH)2SO4	0–80%	Highest BE values obtained with rice bran (10%) and with subsoil as casing material	[43]
P. sajor-caju
P. ostreatus and A. bisporus	NR	P. ostreatus	P. ostreatus and A. bisporus SMS (9:1 and 8:2)		36–40%	Comparable BE values with those obtained using a wheat straw and poultry manure-based commercial substrate	[44]
Pleurotus eous	Wheat straw	P. sajor-caju	SMS with wheat straw (1:9, 1:6 and 1:3)		62–69%	BE increased by increasing SMS content and was higher than the control (wheat straw)	[45]
		P. florida			109 – 113%
		Pleurotus flabellatus			97 – 105%
A. bisporus	Commercial compost with casing	A. bisporus	SMS	Soybean meal (10%), Target® (10%)	26–73%	Lower BE values compared to the conventional substrate	[27]
			SMS and non-composted substrate (1:1 w/w)		97–144%	Similar or higher BE values in comparison to the conventional substrate
Lentinula edodes	Sawdust	P. citrinopileatus	SMS alone and mixed with Mangifera indica sawdust (1:2 w/w)	Rice bran (10%), CaCO3 (pH 6)	23–39%	Increased BE noted when SMS was mixed with M. indica sawdust	[46]
P. sajor-caju					19–34%
P. sajor-caju	Rice straw	P. sajor-caju	SMS	Mustard, niger, sunflower, cotton seed powder and soybean seed cake	45–125%	Better mushroom yields with cotton seed powder than with the other supplements	[47]
F. velutipes	NM	L. edodes	SMS and SMS:oak sawdust (1:4, 2:3, 3:2, 4:1)	Rice bran (20%)	60–84%	High BE values for sawdust ratios above 40%	[48]
A. bisporus	NR	L. edodes	SMS:oak:spruce (1:1:1) and plain SMS	Wheat bran (3%), millet (35%) and rye (2%)	33–53%	Higher BE values for the mixture than for plain SMS	[49]
L. edodes	Sawdust	Pleurotus cornucopiae	SMS:rice bran (1:1)		61%	BE values higher than the control (Abies sachalinensis sawdust) only for P. cornucopiae	[50]
		P. ostreatus	SMS:rice bran (1:1)		62%
		F. velutipes	SMS:wheat bran (1:1)		88%
P. ostreatus	Cryptomeria japonica sawdust	Lyophyllum decastes	SMS and SMS:bark compost (4:1 and 1:1)	Rice bran 10%	62–134%	BE two-fold higher for mixtures plain SMS	[51]
Ph. nameko	Harwood sawdust				140–167%	No significant differences in BE
L. edodes	Oak sawdust	P. sajor-caju	SMS	Soybean flour (0 or 12%), CaCO3 (0 or 1%)	62–79%	Higher BE values with soybean flour supplementation	[52]

Table 2. Reuse of spent mushroom substrate (SMS) as animal feed based on the outcome reported in pertinent publications. Abbreviations used: ADF, acid detergent fiber; ADL, acid detergent lignin; HWE, hot water extract; NDF, neutral detergent fiber; NR, not reported; S-SMS: steam-treated SMS; UN-SMS: untreated SMS.

Origin of SMS	SMS content in fibers and crude protein			Animals in feeding trials	Type of feed, inclusion levels, and length of feeding trials	Main outcome	Ref.
	NFD, ADF and ADL (%)	Crude fiber (%)	Protein (%)
Pleurotus sajor-caju	60.3, 52.3, 4.1	NR	9.3	Alpine dairy goats	Rice straw fermented with P. sajor-caju SMS (5:1 w/w for 8 weeks), diet and water provided ad libitum, 28 days	SMS effectively improved the nutrient content, availability in the rumen, and feeding value of rice straw; increased effective degradability of dry mater and fibers in the rumen; improved intake and milk yield	[61]
Pleurotus ostreatus	NR	NR	3.0	Male Sika deers	SMS (10%), 60 days	SMS led to reduction in the intake of organic matter, and improved digestibility of crude fat	[62]
Flammulina velutipes	NR	NR	5.1			Νo effect on apparent nutrient digestibility, feed intake, velvet antler production, or biochemical indexes
P. ostreatus	NR	NR	NR	Liuyang black goats	SMS co-fermented with feed and whole plant rice, 60 days	Feeding with co-fermented whole plant rice and SMS had no adverse effect on the slaughter performance, while the meat quality was improved	[63]
Cordyceps militaris	NR	NR	NR	Crossbred growing pigs	SMS (0.2%, w/w), 6 weeks	SMS increased final body weight and daily weight gain, immunoglobulin A and G, and the rest of the parameters were not affected.	[55]
Lentinula edodes	NR	NR	NR	Weaned piglets	SMS (3%) fermented by Bacillus subtilis, 33 days	Increase in final weight, daily gain and feed conversion; beneficial effect on the intestinal mucosal barrier, and immunity	[64]
Ganoderma lucidum	NR	NR	23.6	Mice	HWE of SMS (0.14, 0.28, 0.84 and 1.68 g kg^−1), 30 days	HWE of SMS enhanced murine immune function; the 0.84 g/kg dose had optimal effect in all aspects	[65]
P. ostreatus	74.8, 49.4, NR	24.2	7.9	Hanwoo steers	SMS fermented or not with Lactobacillus brevis, 13 days	SMS (fermented or not) could replace formulated feed concentrate without adverse effects	[66]
P. ostreatus	73.6, 55.0, NR	NR	8.1	Geese	SMS fed ad libitum during the growing period, 8 weeks)	SMS supplementation at 5% had no adverse effects on the growth performance, while it favorably affected sensory attributes	[67]
Pleurotus eryngii	50.6*, 38.9*, 11.7*	31.2*	16.9*	Sheep	By-product feed silage with fermented SMS (45%), 22 days	The SMS-containing feed showed similar energy value, higher protein metabolism and utilization, and lower fiber digestion than the rye straw-based control diet	[68]
P. ostreatus	NR	NR	5.6	Awassi sheep	SMS (5, 10, 15 and 20%), 70 days	SMS ratios (>15%) decreased slaughter, empty body, and carcass weights, dressing, leg lean and fat tail percentage, backfat thickness, and rib eye area	[69]
P. ostreatus	76.7, 61.2, 15.5	NR	13.4	Hanwoo steers	Feed containing 50% SMS; ad libitum access during the growing and fattening periods	Trials with by-product feed tended to increase the average daily weight gain and feed efficiency, while it did not affect quality and yield traits	[70]
G. lucidum	NR	6	23.6	Holstein cows	HWE of SMS (33, 67, and 100 g d^−1 cow^−1), 60 days	HWE of SMS may enhance immunity and antioxidant capacity in dairy cows, and subsequently improve milk quality	[71]
Ganoderma chalceum(syn. G. balabacense)	NR	NR	23.6	Chinese Holstein cows	HWE of SMS (33, 67, and 100 g d^−1 cow^−1), 60 days	Feeding by HWE of SMS improved hematology parameters, and increased milk yield, milk protein and triglyceride levels	[72]
Agaricus bisporus	27.8, NR, 6.9	13.3	12.9	Holsteins male calves	SMS (15%), 170 days	No differences detected in the carcass and internal organs of the calves that received different diets	[73]
P. ostreatus	NR	29.6	7.9	Broilers	SMS substituted wheat bran (by 25–100%), 8 weeks	Higher feed intake with increased rate of SMS inclusion; SMS did not affect breast, thigh drumstick, back, neck, wings, and shoulder weight	[74]
Hypsizygus marmoreus	NR	-	-	Laying hens	SMS (5, 10 and 15%) fermented with Bacillus subtilis, 12 weeks	Feed intake increased with SMS addition; no differences in egg production, egg weight, egg mass, feed conversion and viability; the yolk color was more intense when SMS was added	[75]
Grifola frondosa	Steam-treated SMS (S-SMS) and non-treated SMS (N-SMS)	NR	6.6(S-SMS), 6.4 (N-SMS)	Wistar rats	S-SMS or N-SMS (25%), 26 days	Feed with SMS did not affect body weight gain, feed efficiency, or serum biochemical parameters; however, fecal weight and protein content were significantly higher than the control	[76]
P. eryngii	78.8, 66.0. 12.8	NR	5.7	Hanwoo steers during growing and fattening periods	Ad libitum access to microbially-fermented SMS (50%), 12.6 months	Feeding with microbially fermented SMS improved growth performance and carcass traits, and could successfully replace a part of conventional roughage	[77]
P. ostreatus	65.1, 49.4, NR	40.5	15.4	Postweaning calves	SMS (10%) fermented or not with lactic acid bacteria, 60 days	The fermented SMS improved the growth performance compared to non-fermented SMS and to feed with an antibiotic supplement	[78]
P. eryngii	NR	NR	NR	Laying hens	SMS (5, 10 and 15%) fermented with Bacillus subtilis, 7 weeks	Feed intake increased with SMS addition; no differences noted in egg production, egg weight, egg mass, feed conversion and viability; yolk color was more intense when SMS was added	[79]
P. sajor-caju	NR	NR	6.3	Broiler chicken	SMS (0.5 to 2%), 21 or 38 days	The inclusion of SMS up to 0.67% improved the weight gain of broiler chicks in the first 21 days	[80]
Agaricus blazei	NR	NR	NR	Broiler chicks	SMS (0.2 to 1.0%), 42 days	SMS above 0.4% reduced the animals’ performance; 0.2% SMS resulted in the highest weight gain and feed intake, and the best feed conversion	[81]
P. ostreatus	NR	9.4	17.6	Berkshire pigs	SMS (plus rice bran and barley bran, 2:1:1) in ratios of 3, 5 or 7%, 7 weeks	Daily feed intake and feed conversion increased through SMS addition; SMS (3%) positively affected the growth performance, carcass traits, meat quality and fatty acids concentration in meat	[82]
A. bisporus	27.8, 21.0, 6.8	17.8	11.0	Sheep	SMS (10, 20, 30%), 3 weeks	Up to 20% SMS did not affect nutrient intake, digestibility, or nitrogen balance	[83]

*Values referrer to bioproduct-feed with SMS (45%).

Table 3. Reuse of spent mushroom substrate (SMS) as soil amendment based on the outcome reported in pertinent publications. Abbreviations used: NPK, nitrogen, phosphorus, potassium NR, not reported; OC, organic carbon; OM, organic matter; PGPB: plant growth promoting bacteria; TN, total nitrogen.

Origin of SMS	Type of SMS and incorporation rate to soil in field or pot experiments	Effects noted on the soil and/or crops after the use of SMS	Reference
NR	SMS (10–50%) and poultry manure (10–50%) mixed with saline soil in pots	Increase of nutrient availability and salt-tolerant PGPB observed in treated saline soils; using 10% poultry manure and 10% SMS significantly enhanced maize plant growth and yield	[106]
Agaricus bisporus	SMS integrated to the soil in doses of 25 and 100 Mg ha^−1 (dry weight)	SMS in degraded vineyard soils enhanced dehydrogenase activity, respiration activity and soil microbial biomass	[107]
A. bisporus and Pleurotus ostreatus	Composted A. bisporus SMS and P. ostreatus SMS 7:3 (v/v) to replace peat in pots	Higher yields of baby leaf lettuce, i.e. 3–7 times more than that obtained by peat (even under the pressure of the soil-borne plant pathogen Pythium irregulare)	[108]
NR	SMS (20 Mg ha^−1) and chicken manure, applied to sandy soils every 1–2 years for 20 years	OM content increased; pH increased by 1–1.2 units, while soil bulk density decreased; the content of residual pores increased by 30–251%, and the fitted unsaturated hydraulic conductivity decreased	[109]
NR	SMS (35 g) in soil-containing pots (1.5 kg)	Increased NPK and OM contents, soil PGPB, and soil enzyme activities; higher biomass and chlorophyll content obtained in Hibiscus sabdariffa in comparison to the use of mineral NPK (16:16:16) fertilizer	[110]
Flammulina velutipes	Fresh or sterilized SMS (5%, w/w) mixed with soil in glass jars	Total and dissolved OC, microbial biomass carbon and nitrogen, abundance and diversity of bacteria and fungi, and enzyme activities were enhanced	[111]
A. bisporus	SMS (45 and 85 ton ha^−1) mixed with soil in pots	SMS promoted the presence of fungi in the highly connected fraction of the active microbial community	[112]
Auricularia auricula-judae	Composted SMS, biogas residues and pig manure 1:1:1 in seedling pots	Better seedling quality was obtained by using the SMS-based substrate than with the commercial seedling substrates	[113]
Volvariella volvacea	Fresh, weathered, and carbonized SMS mixed with soil (1:2) combined with 0, 50 or 100% of the required rate of nitrogen fertilizer in pots	Weathered and carbonized SMS increased available N; fresh SMS immobilized various nutrients; high yields of pechay during first and second crop on weathered and carbonized SMS; fresh SMS led to high yields only during the third crop; yield was increased by N fertilizer only in weathered and carbonized SMS treatments	[114]
A. bisporus	Composted SMS (5 to 75 g L^−1) in pots	SMS (as the sole fertilizer source) improved grass (Lolium multiflorum) yield up to 300% (with a concentration/dependent response) compared to the untreated control (with no NPK fertilization)	[115]
NR	SMS used to supply 50% or 100% of the crop’s nitrogen requirements	In contrast to mineral fertilizers, no increase in salt content was recorded when SMS was applied; similar lettuce and leek yields when either SMS or mineral fertilizers were used	[116]
P. ostreatus	Fresh SMS incorporated (15–20 t ha^−1) during a period of four years to a depth of approx. 10 cm	SMS led to increase in porosity and fractal dimension, and caused strong development of a granular microstructure in the A horizon (15–20 cm) and a spongy structure in the B horizon (45–50 cm and 70–75 cm)	[117]
A. bisporus, and A. bisporus with P. ostreatus (1:1, v/v)	SMS was incorporated to a soil depth of 30 cm, 1 month prior to planting; both organic treatments providing 100 kg/ha of N	SMS amendment on a calcareous clayey-loam soil resulted in higher oxidizable OC, organic N, extractable K, and available P compared to soil fertilized by 100, 22 and 208 kg/ha N, P and K, respectively; the use of SMS provided lettuce yields similar to that obtained with mineral fertilizer	[105]
NR	SMS and peat moss alone or mixed (1:1, 1:2, and 2:1 (v/v)) with or without NPK fertilizer	SMS could replace up to 50% peat moss to support Chinese kale (Brassica oleracea) production; SMS alone cannot be used as growth medium because of its low nutrient content	[118]
A. bisporus, and A. bisporus with P. ostreatus (1:1, v/v)	SMS-based treatments provided 100 kg ha^−1 of N	SMS increased the oxidizable OC, organic N, available P, respiration rate, and phosphatase activity, while it did not affect pH, EC, catalase, and urease activities in soil cultivated with lettuce	[119]
NR	Fresh or composted SMS applied annually for four years at rates of 8 and 25 Mg ha^−1 (d.w.)	SMS led at increased OC, ΤΝ and labile organic forms as well as enhanced microbiological activity in a semiarid vineyard soil	[120]
Agaricus subrufescens and Lentinula edodes	A. subrufescens SMS (5 to 40%, d.w.) and L. edodes SMS (5 to 25%, d.w.) mixed with soil in pots	SMS led to increase of water retention and enhanced the soil microbial population; when supplemented by 10% of A. subrufescens SMS, lettuce dry weight increased by 2.2 and 1.3 times compared to the control and the NPK (44% N, 37% P2O5 and 48% K2O) treatments; fresh L. edodes SMS did not perform equally well	[104]
NR	SMS distributed onto field plots with a manure spreader at rates of 22.5, 45.0, and 90 kg m^−2	Corn yields were significantly higher in SMS-amended plots, and the nitrogen content of both grain and stover was significantly higher than the control	[121]

Table 4. Reuse of spent mushroom substrate for the control of plant pathogens and pests based on the outcome reported in pertinent publications. Abbreviations used: ACT, aerated compost tea; CT, compost tea; NCT, non-aerated compost tea; NR, not reported.

Origin of SMS	Plant – Pathogen/Pest (disease’s common name)	Main outcome	Reference
Lentinula edodes	Arabidopsis – Alternaria brassicicola	SMS chitin/cellulose nanofiber complex showed disease suppression and growth promotion	[135]
Hypsizygus marmoreus, Pholiota microspora, Lyophyllum decastes, Auricularia polytricha	Arabidopsis – A. brassicicola (cabbage’s leaf spot)	Antifungal volatile compounds emitted by the SMS suppressed fungal infection when incorporated into the soil (1:2, v/v)	[136]
Pleurotus ostreatus	Tomato – Fusarium oxysporum (fusarium wilt)	SMS bio-fortified with Trichoderma asperellum reduced disease severity by 21.2–84.3%	[137]
P. ostreatus, Volvariella volvacea	Pepper – Ralstonia solanacearum, Phytophthora capsica and Meloidogyne spp. (Ralstonia wilt, Phytophthora blight, root-knot nematode)	Biofertilizer (BF) mixed with composted SMS showed a significantly higher disease-control efficacy than BF alone (59 and 76% for P. ostreatus and V. volvacea, respectively, vs. 37% in plain BF)	[138]
P. ostreatus, L. edodes	Tomato – Xanthomonas gardneri (bacterial spot)	Polysaccharides extracted from SMS (1.5 mg mL^−1) reduced bacterial spot severity by 50% on tomato cotyledons, leaflets, and five-leaf plants	[139]
NR	Eggplant – F. oxysporum and R. solanacearum (wilt of eggplant)	SMS, farmyard manure and earthworm compost (1:1:1, w/w) was the most effective combination at inhibiting disease incidence (66.9%)	[140]
L. edodes	Rice – Pyricularia oryzae (rice blast fungus)	Hot-water extract of SMS inhibited the germination of Pyricularia oryzae conidia	[141]
L. edodes	Pepper – Phytophthora capsici (Phytophthora blight)	SMS inhibited mycelial growth of P. capsici, suppressed the Phytophthora blight disease of pepper seedlings by 65% and promoted plant growth by more 30% compared to the control	[142]
Lepista nuda	Cucumber – Pythium aphanidermatum (cucumber’s damping off)	The combination of SMS with peat compost and peat moss reduced the incidence of Pythium damping-off up to 58% and promoted the growth of cucumber seedlings	[143]
Hericium erinaceus	Tomato – R. solanacearum (tomato wilt)	Water extracts of SMS suppressed tomato wilt disease caused by R. solanacearum by 85% in seedlings, and promoted growth of tomato plants	[144]
NR	Tomato – P. capsici (Phytophthora blight)	In vitro bioassays revealed that SMS-ACT reduced P. capsici growth by 50% while SMS-ACT with nutrients reduced it by 66.5%; in greenhouse trials, disease reduction was 6.4–73.4%	[145]
NR	Melon – Didymella bryoniae and Podosphaera fusca (gummy stem blight and powdery mildew)	SMS-ACT and NCT reduced the severity of P. fusca, while only a delay was observed in the growth of D. bryoniae	[146]
L. decastes Pleurotus eryngii	Cucumber – Podosphaera xanthii, Cladosporium cucumerinum, Corynespora cassiicola and Pseudomonas syringae (powdery mildew, cucurbits scab, Corynespora leaf spot and angular leaf spot)	Autoclaved water extract of SMS reduced symptoms caused by P. xanthii and Ps. syringae but not those caused by C. cassiicola and Cl. cucumerinum; a mixture of autoclaved SMS with soil (1:2, v/v), significantly reduced powdery mildew, scab and angular leaf spot diseases	[147]
NR	Bean – F. solani, Rhizoctonia solani and Macrophomina phaseolina (beans root rot)	Soil amendment with SMS-CT was highly effective in reducing root rot incidence caused by F. solani, R. solani and M. phaseolina at pre-emergence damping-off stage and after 45 days	[148]
L. decastes	Cucumber – Colletotrichum lagenarium (anthracnose)	A disease reduction (over 70%) observed in autoclaved and raw SMS incorporated into the soil (1:2, v/v with soil)	[149]
Agaricus bisporus	Tomato – Septoria lycopersici (leaf spot disease)	Plants grown on SMS-containing substrates were resistant to infections caused by S. lycopersici	[150]

Figure 2. Enzymes participating in fungal degradation of lignocellulosic substrates.

Table 5. Recovery of enzymes from spent mushroom substrate. Abbrieviations used: PEG, polyethylene glycol; MnP, manganese peroxidase; LiP, lignin peroxidase; VP, versatile peroxidase.

Mushroom species	Hydrolases	Oxidoreductases	Extraction conditions	Comments	Reference
Agaricus bisporus	Endoxylanase, β-xylosidase, xylan-acetylesterase, arabinofuranosidase, endoglucanase, cellobiohydrolase, β-glucosidase	Peroxidase,phenoloxidase	Distilled water, 0.1 M NaOH, 0.1 M HCl, potassium phosphate buffer, different pH, 37°C, 1 h	No purification; the crude extract was effective for hydrolyzing wheat straw polysaccharides	[166]
A. bisporus		Laccase	Tris – HCl buffer, pH 7.5	No purification; the crude extract was used for oxidation of phenolic compounds	[167]
A. bisporus		Laccase	Distilled water, 4°C, 24 h	Purification by aqueous K3PO4-PEG two-phase system	[168]
A. bisporus	CMCase, xylanase, cellobiohydrolase	VP, MnP, LiP, laccase	Sodium citrate buffer, pH 4.8, room temperature, 150 rpm, 2 h	Acetone precipitation and DEAE chromatography; extract used for hydrolysis of SMS polysaccharides	[169]
Pleurotus sajor-caju	Xylanase, cellulase, β-glucosidase,	Laccase, LiP.	Tap water (pH 8.45), distilled water (pH 5.25), sodium citrate buffer (pH 4.0), 4 or 28°C, 200 rpm, 1–18 h	No purification	[170]
Pleurotus ostreatus, Lentinula edodes, Hericium erinaceus, Flammulina velutipes	α-Amylase, cellulase, xylanase, β-glucosidase	Laccase	1% NaCl, sodium phosphate buffer, tap water, room temperature, shaking, 1 h	No purification	[171]
P. ostreatus, Pleurotus eryngii, Pleurotus cornucopiae	α-Amylase, endoglucanase, endoxylanase	Laccase	Sodium citrate buffer, sodium phosphate buffer, tap water, distilled water, pH 4.5, 4–20°C, 200 rpm, 2–12 h	No purification	[172]
P. eryngii	Xylanase, β-xylosidase, β-glucosidase, α-amylase, cellulase	Laccase, LiP	Tap water, 1% NaOH, phosphate buffer, 4°C, 200 rpm, 2 h	No purification	[173]
P. ostreatus		MnP, laccase, LiP	Sodium tartrate buffer, pH 5.2, 22°C, 150 rpm, 2 h	Ultrafiltration; extract used for soil bioremediation	[174]
Pleurotus florida	CMCase, xylanase, cellobiohydrolase	VP, MnP, LiP, laccase	Sodium citrate buffer, pH 4.8, room temperature, 150 rpm, 2 h	Acetone precipitation, DEAE chromatography	[169]
P. florida	Xylanase	Laccase	Sodium citrate buffer, pH 4.8, room temperature, 2 h	Partial purification by ammonium sulfate precipitation and dialysis	[175]
Pleurotus pulmonarius		Laccase, MnP	Sodium citrate buffer, pH 4.8, 30 min	No purification	[163]
P. pulmonarius	Endoglucanase, xylanase, endoglucanase	LiP, laccase	Tap water, pH 4.0, 4°C, 150 rpm, 1 h	Concentration by freeze drying	[176]
P. ostreatus, Pleurotus citrinopileatus, Auricularia auricula-judae, Coprinus comatus, Agrocybe cylindracea, He. erinaceus, Hypsizygus marmoreus, Tremella fuciformis	Xylanase		Tap water, 25°C, 150 rpm, 3 h	No purification	[177]
L. edodes, He. erinaceus, Stropharia rugosoannulata, Fomes fomentarius, Grifola frondosa	Xylanase	Laccase	Water, 18–48 h, 10 or 20°C	Purification by ultrafiltration, stabilization of the 10 kDa retentate with either glycerol or maltodextrin/sodium benzoate	[178]
L. edodes, P. ostreatus, P. eryngii, Pleurotus spp., F. velutipes, Hypsizygus marmoreus	Total cellulase, CMCase, avicelase, β-glucosidase, dextranase, amylase	Laccase	Distilled water, 30°C, 180 rpm, 1 h	No purification	[179]
Tremella fuciformis	Xylanase, cellulase, pectinase		Tap water, 25°C, 150 rpm, 3 h	Ammonium sulfate precipitation, dialysis, DEAE chromatography	[177]
Ganoderma lucidum		Laccase	Sodium acetate buffer (pH 5.0), liquid-solid ratio 5, 4°C, 3 h	Partial purification by ammonium sulfate precipitation and dialysis	[180]

Table 6. Production of enzymes by fungal cultivation on spent mushroom substrate. Abbreviations used: SSF, solid-state fermentation; SmF, submerged fermentation.

Mushroom species	Enzyme-producing organism	Produced enzymes	Comments	Reference
Auricularia polytricha, Pleurotus ostreatus, Auricularia nigricans	Trichoderma reesei	Cellulase	Ultrasonic-assisted fermentation	[188]
Agaricus bisporus	Trichoderma spp., Aspergillus niger	Endoglucanase, endoxylanase, amylase, β-glucosidase	SSF, no nutritional supplements were used	[189]
Pleurotus pulmonarius	Trichoderma asperellum	1,4-β-cellobiohydrolase,carboxymethylcellulase, β-glucosidase	SSF	[163]
Pleurotus florida	Trichoderma longibrachiatum	Endoglucanase, exoglucanase, xylanase	SSF, SmF	[175]
Pleurotus sajor-caju	Penicillium echinulatum	Endoglucanase, β-glucosidase, xylanase	SmF	[190]
P. florida	Aspergillus aculeatus	Cellobiase	SSF, SmF	[175]
P. ostreatus	P. ostreatus, P. pulmonarius, Ganoderma adspersum, Ganoderma resinaceum, Lentinula edodes	Laccase	SSF of SMS supplemented with wheat bran and soybean flour	[191]
P. ostreatus	P. ostreatus, P. pulmonarius	Laccase	SSF of SMS enriched with wheat bran and soybean flour; the crude enzyme was used for dephenolization of wastewaters	[26]

Table 7. Recovery of bioactive compounds from spent mushroom substrate. Abbreviations used: UAE, ultrasound-assisted extraction; SSF, solid-state fermentation.

Mushroom species	Chemical	Production method	Comment	Reference
Lentinula edodes	Polysaccharide composed of anhydrorhamnose, anhydroglucose, and anhydromannose	Water extraction (90°C, 1 h), alcohol precipitation, centrifugation, freeze-drying, Sevag deproteinization and gel-filtration chromatography	Heteropolysaccharide with antibacterial activity	[195]
L. edodes	Partially hydrolyzed polysaccharides	Water extraction (85°C, 3 h), alcohol precipitation, centrifugation, hydrolysis with 1 M H2SO4, anion-exchange and gel-filtration chromatography	Antioxidant, anti-inflammatory, and renoprotective effects	[57,196]
Ganoderma lucidum	Polysaccharides	Water extraction, Sevag deproteinization and gel-filtration chromatography	Antioxidant activity	[197]
Panus strigellus (syn. Pleurotus tubarius), Pycnoporus sanguineus	β-Glucans, steroids, saturated triterpenes	Extraction with deionized water at 100°C	The extracts were concentrated by lyophilization before analyses	[198]
Pleurotus eryngii	Polysaccharide-protein complex containing 99% carbohydrate and 1% protein	Extraction with 0.5 M NaOH (90°C, 300 min), ethanol precipitation, dialysis, Sevag deproteinization, dialysis, and gel-filtration chromatography	Strong antioxidant activity	[199]
Cordyceps militaris	Polysaccharides	Partial enzymatic hydrolysis (45°C, pH 4.0, 2 h), gradient ethanol precipitation, Sevag deproteinization	Antioxidant activity with no cytotoxicity	[200]
Agrocybe cylindracea, L. edodes, Hypsizygus marmoreus, Pleurotus ostreatus, C. militaris	Polysaccharides	Enzymatic extraction, ethanol precipitation, Sevag deproteinization	Antioxidant activity	[201]
P. ostreatus	Crude exopolysaccharides	SSF of P. ostreatus and P. pulmonarius, EPS extracted with water at 60°C, 15 min	SSF produced laccase, fruitbodies and crude EPS	[191]
L. edodes	Acid polysaccharides	Extraction with .5 M KOH (90°C, 300 min), ethanol precipitation, Sevag deproteinization, purification by gel-filtration chromatography	Heteropolysaccharides with antiproliferative activity against human tumor cells	[193]
P. ostreatus	Crude polysaccharides	Autoclaving with distilled water (120°C, 30 min), ethanol precipitation	Protective effect against plant diseases	[139]
L. edodes	Ergosterol, ergosta-7,22-dienol, β-sitosterol	UAE, optimal conditions: 432 W, 16 min, liquid-solid ratio 22	Antitumor activity	[58]
H. marmoreus	Pentostatin	SSF by C. militaris	Pentostatin is a powerful anticancer drug	[202]

UAE, ultrasound-assisted extraction; SSF, solid-state fermentation.

Figure 3. Schematic illustration of the cascade-use concept of bioresources (based on Vis et al. 2016 [204], p. 6, https://data.europa.eu/doi/10.2873/827106).

Figure 4. Schematic illustration of the concept food – ethanol – solid fuel. The figure is modified from Chen, 2021 [208], p. 15, https://pub.epsilon.slu.se/26324/1/chen_f_211216.pdf.

Ahmad Zakil F, Mohd Isa R, Mohd Sueb MS, et al. Growth performance and mineral analysis of Pleurotus ostreatus (oyster mushroom) cultivated on spent mushroom medium mixed with rubber tree sawdust. Mater Today Proc. 2022;57:1329–1337. doi: 10.1016/j.matpr.2022.01.112

 Google Scholar

Lisiecka J, Prasad R, Jasinska A. The utilisation of Pholiota nameko, Hypsizygus marmoreus, and Hericium erinaceus spent mushroom substrates in Pleurotus ostreatus cultivation. Horticulturae. 2021;7(10):396. doi: 10.3390/horticulturae7100396

 Web of Science ®Google Scholar

Economou CN, Philippoussis AN, Diamantopoulou PA. Spent mushroom substrate for a second cultivation cycle of Pleurotus mushrooms and dephenolization of agro-industrial wastewaters. FEMS Microbiol Lett. 2020;367(8). doi: 10.1093/femsle/fnaa060

 PubMed Web of Science ®Google Scholar

Wu CY, Liang CH, Liang ZC. Evaluation of using spent mushroom sawdust wastes for cultivation of Auricularia polytricha. Agronomy. 2020;10(12):1892. doi: 10.3390/agronomy10121892

 Google Scholar

Lin Q, Long L, Wu L, et al. Evaluation of different agricultural wastes for the production of fruiting bodies and bioactive compounds by medicinal mushroom Cordyceps militaris. J Sci Food Agric. 2017;97(10):3476–3480. doi: 10.1002/jsfa.8097

 PubMed Web of Science ®Google Scholar

Zeng XL, Han F, Ye JL, et al. Recycling spent Pleurotus eryngii substrate supplemented with Tenebrio molitor feces for cultivation of Agrocybe chaxingu. Int J Recycl Org Waste Agricult. 2017;6(4):275–280. doi: 10.1007/s40093-017-0171-9

 Google Scholar

Noonsong V. Recycling of spent Pleurotus compost for production of the Agrocybe cylindracea. Mycosphere. 2016;7(1):36–43. doi: 10.5943/mycosphere/7/1/4

 Web of Science ®Google Scholar

Picornell-Buendía R, Pardo-Giménez A, de Juan-Valero JA. Agronomic assessment of spent substrates for mushroom cultivation. Biotechnol Agron Soc Environ. 2016;20(3):263–374. doi: 10.25518/1780-4507.13138

 Web of Science ®Google Scholar

Picornell MR, Pardo A, de Juan JA. Reuse of degraded Pleurotus ostreatus substrate through supplementation with wheat bran and Calprozime® quantitative parameters. Agron Colomb. 2015;33(2):261–270. doi: 10.15446/agron.colomb.v33n2.49760

 Google Scholar

Wang S, Xu F, Li Z, et al. The spent mushroom substrates of Hypsizigus marmoreus can be an effective component for growing the oyster mushroom Pleurotus ostreatus. Sci Hortic. 2015;186:217–222. doi: 10.1016/j.scienta.2015.02.028

 Web of Science ®Google Scholar

Ashrafi R, Mian MH, Rahman MM, et al. Recycling of spent mushroom substrate for the production of oyster mushroom. Res Biotechnol. 2014;5:37–41.

 Google Scholar

Mishra KK, Pal RS, Arunkumar R, et al. Antioxidant properties of different edible mushroom species and increased bioconversion efficiency of Pleurotus eryngii using locally available casing materials. Food Chem. 2013;138(2–3):1557–1563. doi: 10.1016/j.foodchem.2012.12.001

 PubMed Web of Science ®Google Scholar

Sun X, Xu H, Liang H, et al. Chemical composition of spent Pleurotus eryngii mushroom substrate and its reuse for Volvariella volvacea production. Asian J Chem. 2013;25(18):10504–10508. doi: 10.14233/ajchem.2013.15809

 Google Scholar

Zhang W, Niu X, Zhang W, et al. The cultivation of Agaricus bisporus on the spent substrate of Flammulina velutipes. African Journal Of Agricultural Research. 2013;8:4860–4863.

 Google Scholar

González Matute R, Figlas D, Curvetto N. Agaricus blazei production on non-composted substrates based on sunflower seed hulls and spent oyster mushroom substrate. World J Microbiol Biotechnol. 2011;27(6):1331–1339. doi: 10.1007/s11274-010-0582-5

 PubMed Web of Science ®Google Scholar

Pardo-Giménez A, Pardo-González JE, Zied DC. Evaluation of harvested mushrooms and viability of Agaricus bisporus growth using casing materials made from spent mushroom substrate. Int J Food Sci Technol. 2011;46(4):787–792. doi: 10.1111/j.1365-2621.2011.02551.x

 Web of Science ®Google Scholar

Gern RMM, Libardi Junior N, Patrício GN, et al. Cultivation of Agaricus blazei on Pleurotus spp. spent substrate. Braz Arch Biol Technol. 2010;53(4):939–944. doi: 10.1590/S1516-89132010000400024

 Web of Science ®Google Scholar

Pardo-Giménez A, Pardo-González JE. Elaboration of new substrates for cultivating Pleurotus ostreatus (Jacq.) P. Kumm. based on degraded substrates from edible fungi cultivation. ITEA. 2009;105(2):89–98.

 Google Scholar

Singh C, Oys S. Recycling of spent oyster mushroom substrate to recover additional value. KUSET. 2009;5.

 Google Scholar

Mamiro DP, Royse DJ, Beelman RB. Yield, size, and mushroom solids content of Agaricus bisporus produced on non-composted substrate and spent mushroom compost. World J Microbiol Biotechnol. 2007;23(9):1289–1296. doi: 10.1007/s11274-007-9364-0

 Web of Science ®Google Scholar

Liang ZC, Wu C, Wang J. The evaluation of using mushroom sawdust wastes for cultivation of Pleurotus citrinopileatus. Fungal Sci. 2005;20:27–34.

 Google Scholar

Shashirekha MN, Rajarathnam S, Bano Z. Enhancement of bioconversion efficiency and chemistry of the mushroom, Pleurotus sajor-caju (Berk and br.) Sacc. produced on spent rice straw substrate, supplemented with oil seed cakes. Food Chem. 2002;76(1):27–31. doi: 10.1016/S0308-8146(01)00244-8

 Web of Science ®Google Scholar

Chai J, Kim Y, Lee S. Reutilization of enokitake cultural waste as cultivating substrates for production of shiitake, Lentinus edodes and enokitake, Flammulina velutipes. Gwacheon, South Korea: Report of the Ministry of Agriculture & Forestry of the Republic of Korea; 2000. p. 113.

 Google Scholar

Kilpatrick M, Murray DJ, Ward F Influence of substrate formulation and autoclave treatment on Lentinula edodes production. Proceedings of the 15th International Congress on the Science and Cultivation of Edible Fungi. Maastricht, Netherlands; 2000. p. 803–810.

 Google Scholar

Nakaya M, Yoneyama S, Kato Y, et al. Cultivation of some important edible mushrooms using the sawdust from waste shiitake bed logs. Proceedings of the 3rd International Conference of Mushroom Biology and Mushroom Products. Sydney, Australia: World Society for Mushroom Biology and Mushroom Products; 1999 [cited 2022 May 20]. Available from:https://www.fungifun.org/docs/mushrooms/Shiitake/Mushworld%20Cultivation%20of%20some%20important%20edible%20mushrooms%20using%20the%20sawdust%20from%20waste%20Shiitake%20bed%20logs.pdf

 Google Scholar

Akamatsu Y. Reutilization of culture wastes of Pleurotus ostreatus and Phollota nameko for cultivation of Lyophyllum decastes. J Wood Sci. 1998;44(5):417–420. doi: 10.1007/BF01130458

 Web of Science ®Google Scholar

Royse DJ. Recycling of spent shiitake substrate for production of the oyster mushroom, Pleurotus sajor-caju. Appl Microbiol Biotechnol. 1992;38(2):179–182. doi: 10.1007/BF00174464

 Web of Science ®Google Scholar

Fan GJ, Chen MH, Lee CF, et al. Effects of rice straw fermented with spent Pleurotus sajor-caju mushroom substrates on milking performance in Alpine dairy goats. Anim Biosci. 2022;35(7):999–1009. doi: 10.5713/ab.21.0340

 PubMedGoogle Scholar

Yuan C, Wu M, Tahir SM, et al. Velvet antler production and hematological changes in male Sika deers fed with spent mushroom substrate. Animals (Basel). 2022;12(13):1689. doi: 10.3390/ani12131689

 PubMedGoogle Scholar

Huang LQ, Li SQ, Yuan ZZ, et al. Effects of feeding co-fermented whole plant rice and spent mushroom(Pleurotus ostreatus) substrate on slaughter performance, meat quality and organ size indexes of Liuyang black goats. Acta Prataculturae Sin. 2021;30(6):133.

 Google Scholar

Boontiam W, Wachirapakorn C, Wattanachai S. Growth performance and hematological changes in growing pigs treated with Cordyceps militaris spent mushroom substrate. Vet World. 2019;13(4):768–773. doi: 10.14202/vetworld.2020.768-773

 PubMed Web of Science ®Google Scholar

Qi Q, Peng Q, Tang M, et al. Microbiome analysis investigating the impacts of fermented spent mushroom substrates on the composition of microbiota in weaned piglets hindgut. Front Vet Sci. 2020;7:584243. doi: 10.3389/fvets.2020.584243.

 PubMed Web of Science ®Google Scholar

Liu Y, Mei L, Li J, et al. Effect of JUNCAO-cultivated Ganoderma lucidum spent mushroom substrate-hot water extract on immune function in mice. Trop J Pharm Res. 2018;17(2):261–267. doi: 10.4314/tjpr.v17i2.10

 Web of Science ®Google Scholar

Baek YC, Kim MS, Reddy KE, et al. Rumen fermentation and digestibility of spent mushroom (Pleurotus ostreatus) substrate inoculated with Lactobacillus brevis for Hanwoo steers ¤. Revista Colombiana de Ciencias Pecuarias. 2017;30(4):267–277. doi: 10.17533/udea.rccp.v30n4a02

 Web of Science ®Google Scholar

Chang S, Lin MJ, Chao YP, et al. Effects of spent mushroom compost meal on growth performance and meat characteristics of grower geese. R Bras Zootec. 2016;45(6):281–287. doi: 10.1590/S1806-92902016000600001

 Web of Science ®Google Scholar

Seok JS, Kim YI, Lee YH, et al. Effect of feeding a by-product feed-based silage on nutrients intake, apparent digestibility, and nitrogen balance in sheep. J Anim Sci Technol. 2016;58(1):9. doi: 10.1186/s40781-016-0091-7

 PubMedGoogle Scholar

Aldoori ZT, Al-Obaidi ASA, Abdulkareem AH, et al. Effect of dietary replacement of barley with mushroom cultivation on carcass characteristics of Awassi lambs. J Anim Health Prod. 2015;3(4):94–98. doi: 10.14737/journal.jahp/2015/3.4.94.98

 Google Scholar

Kim YI, Park JM, Lee YH, et al. Effect of by-product feed-based silage feeding on the performance, blood metabolites, and carcass characteristics of Hanwoo steers (a field study). Asian-Australas J Anim Sci. 2015;28(2):180–187. doi: 10.5713/ajas.14.0443

 PubMed Web of Science ®Google Scholar

Liu Y, Zhao C, Lin D, et al. Effects of Ganoderma lucidum spent mushroom substrate extract on milk and serum immunoglobulin levels and serum antioxidant capacity of dairy cows. Trop J Pharm Res. 2015;14(6):1049–1055. doi: 10.4314/tjpr.v14i6.16

 Web of Science ®Google Scholar

Liu Y, Zhao C, Lin D, et al. Effect of water extract from spent mushroom substrate after Ganoderma balabacense cultivation by using JUNCAO technique on production performance and hematology parameters of dairy cows. Anim Sci J. 2015;86(9):855–862. doi: 10.1111/asj.12371

 PubMed Web of Science ®Google Scholar

Fazaeli H, Shafyee-Varzeneh H, Farahpoor A, et al. Recycling of mushroom compost wheat straw in the diet of feedlot calves with two physical forms. Int J Recycl Org Waste Agricult. 2014;3(3):3. doi: 10.1007/s40093-014-0065-z

 Google Scholar

Foluke A, Olutayo A, Olufemi A. Assessing spent mushroom substrate as a replacement to wheat bran in the diet of broilers. Am Int J Contemp Res. 2014;4(4):178–183.

 Google Scholar

Kim SC, Moon YH, Kim HS, et al. Effect of dietary fermented spent mushroom (Hypsizygus marmoreus) substrates on laying hens. J Mushroom. 2014;12(4):350–356. doi: 10.14480/JM.2014.12.4.350

 Google Scholar

Tasaki Y, Kozuka K, Mochida K, et al. Effect of sawdust-based spent mushroom substrate treated with steam on rat growth performance. Food Sci Technol Res. 2014;20(2):493–497. doi: 10.3136/fstr.20.493

 Web of Science ®Google Scholar

Kim YI, Lee YH, Kim KH, et al. Effects of supplementing microbially-fermented spent mushroom substrates on growth performance and carcass characteristics of Hanwoo steers (a field study). Asian-Australas J Anim Sci. 2012;25(11):1575–1581. doi: 10.5713/ajas.2012.12251

 PubMed Web of Science ®Google Scholar

Kim MK, Lee HG, Park JA, et al. Recycling of fermented sawdust-based oyster mushroom spent substrate as a feed supplement for postweaning calves. Asian-Australas J Anim Sci. 2011;24(4):493–499. doi: 10.5713/ajas.2011.10333

 Web of Science ®Google Scholar

Cho YJ, Choi EJ, Shin PG, et al. Effect of spent mushroom (Pleurotus eryngii) substrates addition on egg quality in laying hens. J Mushrooms. 2010;8(4):196–197.

 Google Scholar

Azevedo RS, Ávila CDS, Dias ES, et al. Utilization of the spent substrate of Pleurotus sajor caju mushroom in broiler chicks ration and the effect on broiler chicken performance. Acta Sci - Anim Sci. 2009;31(2):139–144.

 Google Scholar

Machado AMB, Dias ES, dos Santos ÉC, et al. Composto exaurido do cogumelo Agaricus blazei na dieta de frangos de corte. R Bras Zootec. 2007;36(4 suppl):1113–1118. doi: 10.1590/S1516-35982007000500018

 Google Scholar

Song YM, Lee SD, Chowdappa R, et al. Effects of fermented oyster mushroom (Pleurotus ostreats) by-product supplementation on growth performance, blood parameters and meat quality in finishing Berkshire pigs. Animal. 2007;1(2):301–307. doi: 10.1017/S1751731107683785

 PubMed Web of Science ®Google Scholar

Fazaeli H, Masoodi ART. Spent wheat straw compost of Agaricus bisporus mushroom as ruminant feed. Asian-Australas J Anim Sci. 2006;19(6):845–851. doi: 10.5713/ajas.2006.845

 Web of Science ®Google Scholar

Upadhyay SK, Chauhan PK. Optimization of eco-friendly amendments as sustainable asset for salt-tolerant plant growth-promoting bacteria mediated maize (Zea mays L.) plant growth, Na uptake reduction and saline soil restoration. Environ Res. 2022;211:113081. doi: 10.1016/j.envres.2022.113081

 PubMed Web of Science ®Google Scholar

Herrero-Hernández E, Andrades MS, Villalba Eguren G, et al. Organic amendment for the recovery of vineyard soils: effects of a single application on soil properties over two years. Processes. 2022;10(2):317. doi: 10.3390/pr10020317

 Web of Science ®Google Scholar

Hernández D, Ros M, Carmona F, et al. Composting spent mushroom substrate from Agaricus bisporus and Pleurotus ostreatus production as a growing media component for baby leaf lettuce cultivation under Pythium irregulare biotic stress. Horticulturae. 2021;7(2):13. doi: 10.3390/horticulturae7020013

 Web of Science ®Google Scholar

Lipiec J, Usowicz B, Kłopotek J, et al. Effects of application of recycled chicken manure and spent mushroom substrate on organic matter, acidity, and hydraulic properties of sandy soils. Materials. 2021;14(14):4036. doi: 10.3390/ma14144036

 PubMed Web of Science ®Google Scholar

Ngan NM, Riddech N. Use of spent mushroom substrate as an inoculant carrier and an organic fertilizer and their impacts on Roselle growth (Hibiscus sabdariffa L.) and soil quality. Waste Biomass Valor. 2021;12(7):3801–3811. doi: 10.1007/s12649-020-01278-w

 Web of Science ®Google Scholar

Wang HW, Xu M, Cai XY, et al. Evaluation of soil microbial communities and enzyme activities in cucumber continuous cropping soil treated with spent mushroom (Flammulina velutipes) substrate. J Soils Sediments. 2021;21(8):2938–2951. doi: 10.1007/s11368-021-02989-w

 Web of Science ®Google Scholar

Paula FS, Tatti E, Thorn C, et al. Soil prokaryotic community resilience, fungal colonisation and increased cross-domain co-occurrence in response to a plant-growth enhancing organic amendment. Soil Biol Biochem. 2020;149:107937. doi: 10.1016/j.soilbio.2020.107937

 Web of Science ®Google Scholar

Meng X, Liu B, Zhang H, et al. Co-composting of the biogas residues and spent mushroom substrate: Physicochemical properties and maturity assessment. Bioresour Technol. 2019;276:281–287. doi: 10.1016/j.biortech.2018.12.097

 PubMed Web of Science ®Google Scholar

Ultra VU, Sotto JO, Punzalan MR. Nutrient supplying potential of different spent mushroom substrate preparations as soil amendment in a potting media. IOP Conf Ser Earth Environ Sci. 2018;120:012010. doi: 10.1088/1755-1315/120/1/012010

 Google Scholar

Paula FS, Tatti E, Abram F, et al. Stabilisation of spent mushroom substrate for application as a plant growth-promoting organic amendment. J Environ Manage. 2017;196:476–486. doi: 10.1016/j.jenvman.2017.03.038

 PubMed Web of Science ®Google Scholar

Gobbi V, Bonato S, Nicoletto C, et al. Spent mushroom substrate as organic fertilizer: vegetable organic trials | International society for horticultural science. Proceedings of the III International Symposium on Organic Matter Management and Compost Use in Horticulture. Murcia, Spain; 2015. p. 49–56.

 Google Scholar

Nakatsuka H, Oda M, Hayashi Y, et al. Effects of fresh spent mushroom substrate of Pleurotus ostreatus on soil micromorphology in Brazil. Geoderma. 2016;269:54–60. doi: 10.1016/j.geoderma.2016.01.023

 Web of Science ®Google Scholar

Paredes C, Medina E, Bustamante MA, et al. Effects of spent mushroom substrates and inorganic fertilizer on the characteristics of a calcareous clayey-loam soil and lettuce production. Soil Use And Manag. 2016;32(4):487–494. doi: 10.1111/sum.12304

 Web of Science ®Google Scholar

Sendi H, Mohamed MTM, Anwar MP, et al. Spent mushroom waste as a media replacement for peat moss in Kai-Lan (Brassica oleracea var. alboglabra) production. Sci World J. 2013;2013:1–8. doi: 10.1155/2013/258562

 Google Scholar

Medina E, Paredes C, Bustamante MA, et al. Relationships between soil physico-chemical, chemical and biological properties in a soil amended with spent mushroom substrate. Geoderma. 2012;173–174:152–161. doi: 10.1016/j.geoderma.2011.12.011

 Web of Science ®Google Scholar

Peregrina F, Larrieta C, Colina M, et al. Spent mushroom substrates influence soil quality and nitrogen availability in a semiarid vineyard soil. Soil Sci Soc Am J. 2012;76(5):1655–1666. doi: 10.2136/sssaj2012.0018

 Web of Science ®Google Scholar

Ribas LCC, de Mendonça MM, Camelini CM, et al. Use of spent mushroom substrates from Agaricus subrufescens (syn. A. blazei, A. brasiliensis) and Lentinula edodes productions in the enrichment of a soil-based potting media for lettuce (Lactuca sativa) cultivation: Growth promotion and soil bioremediation. Bioresour Technol. 2009;100(20):4750–4757.

 PubMed Web of Science ®Google Scholar

Wuest PJ, Fahy HK, Fahy J. Use of spent mushroom substrate (SMS) for corn (maize) production and its effect on surface water quality. Compost Sci Util. 1995;3(1):46–54. doi: 10.1080/1065657X.1995.10701768

 Web of Science ®Google Scholar

Li H, Yoshida S, Mitani N, et al. Disease resistance and growth promotion activities of chitin/cellulose nanofiber from spent mushroom substrate to plant. Carbohydr Polym. 2022;284:119233. doi: 10.1016/j.carbpol.2022.119233

 PubMed Web of Science ®Google Scholar

Fujita R, Yokono M, Ube N, et al. Suppression of Alternaria brassicicola infection by volatile compounds from spent mushroom substrates. J Biosci Bioeng. 2021;132(1):25–32. doi: 10.1016/j.jbiosc.2021.03.003

 PubMed Web of Science ®Google Scholar

Singh G, Tiwari A, Gupta A, et al. Bioformulation development via valorizing silica-rich spent mushroom substrate with Trichoderma asperellum for plant nutrient and disease management. J Environ Manage. 2021;297:113278. doi: 10.1016/j.jenvman.2021.113278

 PubMed Web of Science ®Google Scholar

Yu YY, Li SM, Qiu JP, et al. Combination of agricultural waste compost and biofertilizer improves yield and enhances the sustainability of a pepper field. J Plant Nutr Soil Sci. 2019;182(4):560–569. doi: 10.1002/jpln.201800223

 Google Scholar

Aguiar T, Luiz C, Neto A, et al. Residual polysaccharides from fungi reduce the bacterial spot in tomato plants. Bragantia. 2018;77(2):299–313. doi: 10.1590/1678-4499.2016514

 Web of Science ®Google Scholar

Biswas MK, Ghosh T. Screening of brinjal genotypes for their resistance against fungal and bacterial wilt and integrated management of the disease. Plant Cell Biotechnol Mol Biol. 2018;19:61–71.

 Google Scholar

Ishihara A, Goto N, Kikkawa M, et al. Identification of antifungal compounds in the spent mushroom substrate of Lentinula edodes. J Pestic Sci. 2018;43(2):108–113.

 PubMed Web of Science ®Google Scholar

Kang DS, Min KJ, Kwak AM, et al. Defense response and suppression of phytophthora blight disease of pepper by water extract from spent mushroom substrate of Lentinula edodes. Plant Pathol J. 2017;33(3):264–275. doi: 10.5423/PPJ.OA.02.2017.0030

 PubMed Web of Science ®Google Scholar

Chen JT, Lin MJ, Huang JW. Efficacy of spent blewit mushroom compost and Bacillus aryabhattai combination on control of Pythium damping-off in cucumber. J Agric Sci. 2015;153(7):1257–1266. doi: 10.1017/S0021859614000987

 Web of Science ®Google Scholar

Kwak AM, Min KJ, Lee SY, et al. Water extract from spent mushroom substrate of Hericium erinaceus suppresses bacterial wilt disease of tomato. Mycobiology. 2015;43(3):311–318. doi: 10.5941/MYCO.2015.43.3.311

 PubMed Web of Science ®Google Scholar

Nicol RW, Burlakoti P. Effect of aerobic compost tea inputs and application methods on protecting tomato from Phytophthora capsici. Acta Hortic. 2015;1069(1069):229–233. doi: 10.17660/ActaHortic.2015.1069.32

 Google Scholar

Marín F, Santos M, Diánez F, et al. Characters of compost teas from different sources and their suppressive effect on fungal phytopathogens. World J Microbiol Biotechnol. 2013;29(8):1371–1382. doi: 10.1007/s11274-013-1300-x

 PubMed Web of Science ®Google Scholar

Parada RY, Murakami S, Shimomura N, et al. Suppression of fungal and bacterial diseases of cucumber plants by using the spent mushroom substrate of Lyophyllum decastes and Pleurotus eryngii. J Phytopathol. 2012;160(7–8):390–396. doi: 10.1111/j.1439-0434.2012.01916.x

 Web of Science ®Google Scholar

Mansour F, El-Sayed G. Soil amendment and seed treatments with compost tea as alternative fungicide for controlling root rot disease of bean plants - proquest. Egypt J Biol Pest Control. 2011;21(1):19–26.

 Web of Science ®Google Scholar

Parada RY, Murakami S, Shimomura N, et al. Autoclaved spent substrate of hatakeshimeji mushroom (Lyophyllum decastes Sing.) and its water extract protect cucumber from anthracnose. Crop Prot. 2011;30(4):443–450. doi: 10.1016/j.cropro.2010.11.021

 Web of Science ®Google Scholar

Ntougias S, Papadopoulou KK, Zervakis GI, et al. Suppression of soil-borne pathogens of tomato by composts derived from agro-industrial wastes abundant in mediterranean regions. Biol Fertil Soils. 2008;44(8):1081–1090. doi: 10.1007/s00374-008-0295-1

 Web of Science ®Google Scholar

Ball AS, Jackson AM. The recovery of lignocellulose-degrading enzymes from spent mushroom compost. Bioresour Technol. 1995;54(3):311–314. doi: 10.1016/0960-8524(95)00153-0

 Web of Science ®Google Scholar

Trejo-Hernandez MR, Lopez-Munguia A, Ramirez RQ. Residual compost of Agaricus bisporus as a source of crude laccase for enzymic oxidation of phenolic compounds. Process Biochem. 2001;36(7):635–639. doi: 10.1016/S0032-9592(00)00257-0

 Web of Science ®Google Scholar

Mayolo-Deloisa K, Del Refugio Trejo-Hernandez M, Rito-Palomares M. Recovery of laccase from the residual compost of Agaricus bisporus in aqueous two-phase systems. Process Biochemistry query. 2009;44(4):435–439. doi: 10.1016/j.procbio.2008.12.010

 Web of Science ®Google Scholar

Devi R, Kapoor S, Thakur R, et al. Lignocellulolytic enzymes and bioethanol production from spent biomass of edible mushrooms using Saccharomyces cerevisiae and Pachysolen tannophilus. Biomass Convers Biorefin. 2022;1–15. doi: 10.1007/s13399-022-02406-3

 Web of Science ®Google Scholar

Singh AD, Abdullah N, Vikineswary S. Optimization of extraction of bulk enzymes from spent mushroom compost. J Chem Technol Biotechnol. 2003;78(7):743–752. doi: 10.1002/jctb.852

 Web of Science ®Google Scholar

Ko HG, Park SH, Kim SH, et al. Detection and recovery of hydrolytic enzymes from spent compost of four mushroom species. Folia Microbiol. 2005;50(2):103. doi: 10.1007/BF02931456

 PubMed Web of Science ®Google Scholar

Lim SH, Lee YH, Kang HW. Efficient recovery of lignocellulolytic enzymes of spent mushroom compost from oyster mushrooms, Pleurotus spp., and potential use in dye decolorization. Mycobiology. 2013;41(4):214–220. doi: 10.5941/MYCO.2013.41.4.214

 PubMed Web of Science ®Google Scholar

Lim SH, Kim JK, Lee YH. Production of lignocellulolytic enzymes from spent mushroom compost of Pleurotus eryngii. Korean J Mycol. 2012;40(3):152–158. doi: 10.4489/KJM.2012.40.3.152

 Google Scholar

Sadiq S, Hassan MM, Saba S, et al. Bioremediation of endosulfan in soil using ligninolytic extract of spent mushroom compost of Pleurotus ostreatus. Appl Ecol Environ Res. 2019;17(2):3251–3267. doi: 10.15666/aeer/1702_32513267

 Web of Science ®Google Scholar

Rajavat AS, Rai S, Pandiyan K, et al. Sustainable use of the spent mushroom substrate of Pleurotus florida for production of lignocellulolytic enzymes. J Of Basic Microbiol. 2020;60(2):173–184. doi: 10.1002/jobm.201900382

 PubMed Web of Science ®Google Scholar

Ariff INM, Bahrin EK, Ramli N, et al. Direct use of spent mushroom substrate from Pleurotus pulmonarius as a readily delignified feedstock for cellulase production. Waste Biomass Valori. 2019;10(4):839–850. doi: 10.1007/s12649-017-0106-8

 Web of Science ®Google Scholar

Yunan NAM, Shin TY, Ibrahim S, et al. Upcycling the spent mushroom substrate of the grey oyster mushroom Pleurotus pulmonarius as a source of lignocellulolytic enzymes for palm oil mill effluent hydrolysis. J Microbiol Biotechnol. 2021;31(6):823–832. doi: 10.4014/jmb.2103.03020

 PubMed Web of Science ®Google Scholar

Lin H, Sun M, Li J, et al. Purification and characterization of xylanase from spent mushroom compost and its application in saccharification of biomass wastes. BioResources. 2017;13(1):220–230. doi: 10.15376/biores.13.1.220-230

 Web of Science ®Google Scholar

Schimpf U, Schulz R. Industrial by-products from white-rot fungi production. Part I: Generation of enzyme preparations and chemical, protein biochemical and molecular biological characterization. Process Biochem. 2016;51(12):2034–2046. doi: 10.1016/j.procbio.2016.08.032

 Web of Science ®Google Scholar

Nakajima VM, Soares FEF, Queiroz JH. Screening and decolorizing potential of enzymes from spent mushroom composts of six different mushrooms. Biocatal Agric Biotechnol. 2018;13:58–61. doi: 10.1016/j.bcab.2017.11.011

 Web of Science ®Google Scholar

Liao CS, Yuan SY, Hung BH, et al. Removal of organic toxic chemicals using the spent mushroom compost of Ganoderma lucidum. J Environ Monit. 2012;14(7):1983–1988. doi: 10.1039/c2em10910g

 PubMedGoogle Scholar

He J, Qiu Y, Ji X, et al. A novel strategy for producing cellulase from Trichoderma reesei with ultrasound‐assisted fermentation using spent mushroom substrate. Process Biochem. 2021;104:110–116. doi: 10.1016/j.procbio.2021.03.015

 Web of Science ®Google Scholar

Grujić M, Dojnov B, Potočnik I, et al. Spent mushroom compost as substrate for the production of industrially important hydrolytic enzymes by fungi Trichoderma spp. and Aspergillus niger in solid state fermentation. Int Biodeter Biodegr. 2015;104:290–298. doi: 10.1016/j.ibiod.2015.04.029

 Web of Science ®Google Scholar

Camassola M, Dillon AJP. Biological pretreatment of sugar cane bagasse for the production of cellulases and xylanases by Penicillium echinulatum. Ind Crops Prod. 2009;29(2):642–647. doi: 10.1016/j.indcrop.2008.09.008

 Web of Science ®Google Scholar

Economou CN, Diamantopoulou PA, Philippoussis AN. Valorization of spent oyster mushroom substrate and laccase recovery through successive solid state cultivation of Pleurotus, Ganoderma, and Lentinula strains. Appl Microbiol Biotechnol. 2017;101(12):5213–5222. doi: 10.1007/s00253-017-8251-3

 PubMed Web of Science ®Google Scholar

Zhu H, Sheng K, Yan E, et al. Extraction, purification and antibacterial activities of a polysaccharide from spent mushroom substrate. Int j biol macromol. 2012;50(3):840–843. doi: 10.1016/j.ijbiomac.2011.11.016

 PubMed Web of Science ®Google Scholar

Song X, Ren Z, Wang X, et al. Antioxidant, anti-inflammatory and renoprotective effects of acidic-hydrolytic polysaccharides by spent mushroom compost (Lentinula edodes) on LPS-induced kidney injury. Int j biol macromol. 2020;151:1267–1276. doi: 10.1016/j.ijbiomac.2019.10.173

 PubMed Web of Science ®Google Scholar

Ren Z, Liu W, Song X, et al. Antioxidant and anti-inflammation of enzymatic-hydrolysis residue polysaccharides by Lentinula edodes. Int j biol macromol. 2018;120:811–822. doi: 10.1016/j.ijbiomac.2018.08.114

 PubMed Web of Science ®Google Scholar

Zhang J, Meng G, Zhai G, et al. Extraction, characterization and antioxidant activity of polysaccharides of spent mushroom compost of Ganoderma lucidum. Int j biol macromol. 2016;82:432–439. doi: 10.1016/j.ijbiomac.2015.10.016

 PubMed Web of Science ®Google Scholar

Dávila-Giraldo LR, Zambrano-Forero C, Torres-Arango O, et al. Integral use of rice husks for bioconversion with white-rot fungi. Biomass Convers Biorefin. 2022;12(8):2981–2991. doi: 10.1007/s13399-020-00940-6

 Web of Science ®Google Scholar

He P, Li F, Huang L, et al. Chemical characterization and antioxidant activity of polysaccharide extract from spent mushroom substrate of Pleurotus eryngii. J Taiwan Inst Chem Eng. 2016;69:48–53. doi: 10.1016/j.jtice.2016.10.017

 Web of Science ®Google Scholar

Zeng Y, Zhang Y, Zhang L, et al. Structural characterization and antioxidant and immunomodulation activities of polysaccharides from the spent rice substrate of Cordyceps militaris. Food Sci Biotechnol. 2015;24(5):1591–1596. doi: 10.1007/s10068-015-0205-x

 Web of Science ®Google Scholar

Zhang Y, Zeng Y, Zhang LJ, et al. Monosaccharide composition and antioxidant activity of polysaccharides from different spent mushroom substrates. Food Sci. 2015;36(5):18–23.

 Google Scholar

Zhang Y, Liu W, Huang W, et al. Characterization and antiproliferative effect of novel acid polysaccharides from the spent substrate of shiitake culinary-medicinal mushroom Lentinus edodes (Agaricomycetes) cultivation. Int J Medic Mushrooms. 2017;19(5):395–403. doi: 10.1615/IntJMedMushrooms.v19.i5.20

 PubMed Web of Science ®Google Scholar

Wang Q, Cheng J, Wang L, et al. Valorization of spent shiitake substrate for recovery of antitumor fungal sterols by ultrasound-assisted extraction. J Food Biochem. 2018;42(5):e12602. doi: 10.1111/jfbc.12602

 Web of Science ®Google Scholar

Zou G, Li B, Wang Y, et al. Efficient conversion of spent mushroom substrate into a high value-added anticancer drug pentostatin with engineered Cordyceps militaris. Green Chem. 2021;23(24):10030–10038. doi: 10.1039/D1GC03594K

 Web of Science ®Google Scholar

Vis M, Mantau U, Allen B, Eds. Study on the optimised cascading use of wood Figure 3. https://data.europa.eu/doi/10.2873/827106

 Google Scholar

Chen F Combined production of edible mushrooms and biofuels from lignocellulosic residues [Internet] [ Doctoral thesis]. 2021 [cited 2022 Aug 22]. Available from: https://pub.epsilon.slu.se/26324/1/chen_f_211216.pdf

 Google Scholar

Data availability statement

Since no new data were created by the authors, data sharing is not applicable regarding their own research results. Raw data on search criteria used for building the review, as well as the excluded material, will be available upon reasonable request.