AGRICULTURAL AND FOOD SCIENCE Agricultural and Food Science (2025) 34: xxx–xxx 1 https://doi.org/10.23986/afsci.163895 Spent mushroom substrate (SMS) as a sustainable soil amendment and biofertilizer: A review of opportunities and challenges in agricultural and horticultural systems Ida Quisth1,‡, Isabella Ulfsparre1,‡, Bettina Müller1, Volkmar Passoth1, Svein Øivind Solberg2, Carlos Martín2,3, Luis A. Romero-Soto2, Mari Pent4 and Hesam Mousavi2 1 Department of Molecular Sciences, Swedish University of Agricultural Sciences, Uppsala BioCentrum, Box 7015, SE 75007 Uppsala, Sweden 2 Faculty of Applied Ecology, Agricultural Sciences, and Biotechnology, University of Inland Norway, 2418 Elverum, Norway 3 Department of Chemistry, Umeå University, SE-901 87 Umeå, Sweden 4 Institute of Ecology and Earth Sciences, University of Tartu, 51005 Tartu, Estonia ‡ These authors contributed equally to this work. e-mail: hesam.mousavi@inn.no Spent mushroom substrate (SMS), a byproduct of mushroom cultivation, has gained increasing attention as a sustain- able soil amendment in agricultural and horticultural systems. SMS provides a promising alternative to peat-based substrates and synthetic fertilizers, supporting circular bioeconomy principles and sustainable farming practices. This is particularly relevant in boreal regions, where soils are typically acidic, nutrient-poor, and subject to short growing season conditions, presenting both opportunities and challenges for organic inputs. This review synthesizes current research on the agronomic and environmental implications of SMS use in agriculture and horticulture, including its effects on seed germination, plant growth, crop yield, root development, soil structure, and disease suppression. From an environmental perspective, the use of SMS contributes to waste reduction by repurposing organic residues and replacing peat, a non-renewable resource with significant ecological costs. However, several hurdles remain, including high salinity, inconsistent composition, nutrient imbalances, and complex compounds difficult for plants to access, which can impede plant performance. While prior studies have explored SMS in isolated settings, a comprehensive evaluation across systems is lacking. This review addresses that gap by assessing current evidence, identifying limitations, and outlining future research needs to optimize the use of SMS and scale its adoption in sustainable farming. Key words: sustainable agriculture, peat alternative, organic waste recovery, circular bioeconomy, soil health, environmental safety Introduction Spent mushroom substrate (SMS) is the residual biomass generated after ending the mushroom cultivation cycle in lignocellulose-based substrates (Martín et al. 2023b). Once the mushrooms are harvested, a substantial quan- tity of SMS remains. Although the SMS still contains a part of the carbohydrates and lignin contained in the initial substrate (Klausen et al. 2025), it is depleted of the nutrients needed to support an additional mushroom pro- duction cycle (Kousar et al. 2024). SMS typically consists of partially decomposed lignocellulosic biomass, fungal mycelium remnants, and a small amount of non-utilized nutrients. It is rich in organic compounds, including pro- teins, polysaccharides, lipids, and phenolic compounds—many of which are beneficial for plant growth and soil health (Velusami et al. 2021, He et al. 2024). For each kilogram of fresh mushrooms produced, up to 5 kg of SMS is generated, totaling over 60 million tons of SMS worldwide in 2018 (Zhu et al. 2018, Meshram et al. 2024), and this amount is projected to reach approximately 104 million tons by 2026 (Atallah et al. 2021). Due to its high organic matter content and presence of essential macro- and micronutrients, SMS has gained attention as a sustainable input in both agricultural and horticultural systems. Studies have shown that SMS from various mushroom species can enhance plant growth, act as an organic fertilizer, improve seedling development, and serve as an effective soil amendment (Velusami et al. 2021, Alves et al. 2022). Additionally, SMS harbors diverse microbial communities that not only support plant health but may also suppress soilborne pathogens such as nematodes and fungi. Its potential in bioremediation has also been explored, particularly in its capacity to remediate pollutants and contaminants from soils (Malik et al. 2021). Received 1 September 2025 / Accepted 4 November 2025 The Scientific Agricultural Society of Finland ©This is an open access article under the CC BY 4.0 AGRICULTURAL AND FOOD SCIENCE Agricultural and Food Science (2025) 34: xxx–xxx 2 https://doi.org/10.23986/afsci.163895 SMS can be used in its raw form or processed into spent mushroom compost, a more stable, nutrient-rich product. When incorporated into peat-based growing media or soil in varying proportions, SMS has been shown to improve plant yield, root development, and growth, depending on crop type (Khalil et al. 2024). Moreover, it has been associated with enhanced soil physical properties, including better water retention, increased aggregate stability, and improved filtration capacity (Courtney and Mullen 2009, Priadi and Saskiawan 2018) Environmental concerns about the use of peat—a non-renewable resource—have accelerated interest in alterna- tives such as SMS. Peat, while valued for its physicochemical properties, contributes significantly to greenhouse gas (GHG) emissions and the degradation of wetlands. Several countries have already implemented restrictions or bans on peat use (Gao et al. 2015). Consequently, SMS is being considered a more sustainable alternative, as research indicates it could effectively replace at least parts of growing media (Hernández et al. 2021, Oliveira Vieira et al. 2022). Therefore, integrating SMS into horticultural and agricultural systems provides dual benefits: enhancing crop productivity and promoting circularity by diverting organic waste from landfills. Its valorization supports more resilient, eco-friendly food systems and contributes to global efforts to reduce the environmental impact of farming (Zied et al. 2020). Objectives and scope of review The purpose of this review is to critically assess the emerging use of SMS in agriculture and horticulture, focusing on its agronomic benefits, functionality as a biofertilizer, soil amendment, and peat substitute, and its environ- mental implications. Firstly, the review aims to evaluate the potential of SMS as a biofertilizer and soil amendment by analyzing reported data on its nutrient composition, organic matter content, and effects on soil fertility. Secondly, it seeks to explore the agronomic benefits of SMS by assessing its impact on soil properties, crop productivity, and long- term soil health. This includes investigating improvements in soil structure, nutrient retention, and microbial activity, as well as their influence on plant growth and yield. Finally, the review will explore the environmental implications of SMS use and highlight the advantages and limitations of SMS, compared to traditional biofertilizers and soil amendments, in sustainable farming practices. Collectively, through a thorough analysis of the existing literature, this review aims to provide an understanding of SMS’s viability as a valuable resource in modern farming systems, along with suggestions to guide further development and application. Methodology Systematic literature review This review employs a systematic literature review approach to provide a comprehensive and unbiased assess- ment of existing research on the use of SMS. The approach involves selecting and analyzing relevant studies on the use of SMS as a soil amendment, peat substitute, and biofertilizer in agricultural and horticultural settings, along with their environmental implications. Search strategy A structured search was conducted in the Web of Science and Scopus databases to identify peer-reviewed, high- quality research articles published between 2009 and March 2025, written in English, and available as open access. The primary search terms included: “Spent Mushroom Substrate,” “biofertilizer,” “soil amendment,” “peat substitute,” “growing media,” “horticulture,” “agriculture,” and “environment.” Each keyword was supplemented with relevant free-text variants and Boolean operators to capture different word combinations. The whole search strategy, including exact search strings and combinations, is provided in the Appendix (Tables 1 and 2). In addition to database searches, a manual review of reference lists and citations was conducted to identify further relevant studies. Where access was restricted, article titles or DOIs were used to locate full texts via institutional re- positories or third-party sources. In some cases, EndNote was used to retrieve full-text PDFs directly. Studies were included if they investigated the use of SMS in agriculture or horticulture and explicitly focused on plant growth, soil properties, peat substitution, or environmental effects. Although review articles were not explicitly excluded, the emphasis was on original research, and review articles were given lower priority during the selection process. AGRICULTURAL AND FOOD SCIENCE Agricultural and Food Science (2025) 34: xxx–xxx 3 https://doi.org/10.23986/afsci.163895 Data extraction and analysis From the selected studies, data were extracted on SMS use in agricultural and horticultural settings, including crop type, soil type, application methods, SMS-to-soil ratios, and measured outcomes such as plant growth, yield, or soil improvement. Environmental implications, limitations, and methodological characteristics were also recorded. Key findings from each study were tabulated and grouped by thematic areas (e.g., crop type, application rate, ob- served effect, and method of application). A cross-comparison was then performed to identify recurring trends and patterns, such as how different SMS proportions influence crop performance or soil health. To ensure reli- ability, all included studies were reviewed critically, and extracted data were double-checked to reduce the risk of misinterpretation or omission. SMS as a soil amendment in agriculture A growing body of research has investigated the effects of SMS as a soil amendment in agriculture. This section synthesizes current knowledge on how SMS influences crop productivity, soil properties, microbial activity, and long-term agronomic performance. Table 1 consolidates the agronomic impacts of SMS on crop yields, soil health, and long-term fertility across diverse agricultural contexts. Table 1. Integrated summary of spent mushroom substrate (SMS) applications as a bio-based soil amendment, fertilizer, and peat substitute in agriculture, horticulture, and environmental management Aspect / Category Key Findings Representative References Nutrient Content & Fertility SMS is rich in macronutrients (N, P, K, Ca, Mg) and beneficial microbial biomass; composting enhances nutrient availability and stability. Some micronutrients (e.g., Fe, Zn, Mn) may be lacking. Medina et al. 2009; Postinguel et al. 2025; Domínguez-Gutiérrez et al. 2022; Ravlikovsky et al. 2024; Alves et al. 2022 Crop Productivity Increases growth, biomass, and yield in cereals, legumes, vegetables, and ornamentals; improves germination and seedling vigor; performance varies by SMS type, dose, and processing. He et al. 2024; Elsakhawy et al. 2020a, b; Amarasinghe and Jayaweera 2022; Roy et al. 2015 Nutrient Uptake & Efficiency Increases uptake of P, K, S, nitrate; improves protein content and nutrient cycling; may replace synthetic topdressings. Domínguez-Gutiérrez et al. 2022; Ngan & Riddech 2021; Wiśniewska- Kadżajan and Malinowska 2022 Soil Physical Properties Improves structure, porosity, aeration, water retention, root penetration, and reduces bulk density; enhances soil temperature regulation and aggregation. Ma et al. 2025; Nakatsuka et al. 2016; Kumar and Chugh 2022; Marín-Benito et al. 2016 Soil Chemical Properties Boosts SOC, CEC, P, and K availability, pH buffering; composting stabilizes pH and reduces EC; nutrient cycling is improved. Pan et al. 2023; Igual et al. 2024; Mahato et al. 2024; Gupta et al. 2024 Soil Biological Properties Enhances microbial biomass, respiration, enzymatic activity, and diversity; promotes beneficial microbes (Trichoderma, Pseudomonas); supports pollutant degradation. Martín et al. 2023b; Kwiatkowska et al. 2024; Paula et al. 2020; Prasad et al. 2023 Disease Suppression & Stress Resilience SMS enhances rhizosphere microbiota and suppresses pathogens (e.g., Fusarium), effective in tomato, lettuce, sesame, cucumber, and chili. Improves plant tolerance to drought and salinity; increases shoot/root biomass and physiological resilience when combined with fertilization. Wang et al. 2020; Mahato et al. 2024; Kumar and Chugh 2022; Hernández et al. 2021; Postinguel et al. 2025 Environmental Benefits Substitutes for peat and synthetic fertilizers; reduces GHG emissions and environmental degradation; enhances ecosystem resilience, water retention, and erosion control. Meng et al. 2018; Zakaria et al. 2023; Othman et al. 2020; Roy et al. 2015 Circular Economy & Waste Valorization Converts agro-waste into valuable input; reduces open dumping, transport emissions, and landfill use; integrates into circular agriculture. Martín et al. 2023a; Singh et al. 2021; Thygesen et al. 2021; Kousar et al. 2024 Carbon Sequestration SMS contains slowly decomposing lignocellulosic matter; composting facilitates humification and long-term SOC accumulation; boosts microbial C stabilization. Singh et al. 2021; Leong et al. 2022; Afsar et al. 2024; Khalil et al. 2024 Long-Term Soil Health Sustains fertility through slow nutrient release and organic matter buildup; enhances microbial networks and structural stability over time. Ma et al. 2025; Lou et al. 2017; Paula et al. 2020; Grimm and Wösten 2018 Peat Replacement Comparison Compared to peat, SMS has higher macronutrient levels and microbial activity but less uniformity and lower organic matter; suitable when mixed with compost or leached. Medina et al. 2009; Gao et al. 2015; Hernández et al. 2021; Roy et al. 2015 Best Use Practices Compost or leach SMS before use; blend with peat or compost; enrich with microbes/nutrients; match SMS properties to crop needs; monitor EC/pH regularly. Prasad et al. 2021; Amarasinghe and Jayaweera 2022; Kahn et al. 2012 AGRICULTURAL AND FOOD SCIENCE Agricultural and Food Science (2025) 34: xxx–xxx 4 https://doi.org/10.23986/afsci.163895 SMS effect on agricultural productivity Several studies have demonstrated that SMS can enhance crop productivity through various mechanisms. How- ever, outcomes depend heavily on the quality of SMS, the application method, and the environmental context in which it is used. For instance, Courtney and Mullen (2009) reported a 50% increase in barley (Hordeum vulgare) yield following SMS application, a result comparable to that achieved with inorganic fertilizers. Their field exper- iment assessed different rates of SMS, aeration compost, and synthetic fertilizer on barley, revealing that SMS significantly improved soil organic carbon (SOC), available nutrients, and cation levels without raising heavy met- al concentrations to concerning levels. The strongest correlations between increased soil nutrient content and barley yield were observed in plots with higher SMS application rates, up to 100 t ha-1, indicating SMS’s capacity to improve crop yields. Similarly, Wiśniewska-Kadżajan and Malinowska (2022) studied the combined use of SMS and synthetic N in grass production (Festulolium braunii). They observed increased protein content, biomass formation, and digestibility, with the most effective treatment involving 15 t ha⁻¹ of SMS combined with 68 kg ha⁻¹ of synthetic N, demonstrat- ing SMS’s potential to replace synthetic N inputs partially. In maize (Zea mays), SMS use was associated with improved germination and seedling vigor. In their pot study, Alves et al. 2022 demonstrated that incorporating spent Agaricus subrufescens substrate—especially when com- bined with synthetic fertilization at sowing—led to significant increases in germination rates and pace, as well as other key successive growth parameters, such as shoot height, stem diameter, fresh and dry biomass, and leaf area index. The SMS treatment increased the concentrations of P, K, and S in the biomass and increased soil K, suggesting it could reduce the need for synthetic fertilizers at the topdressing stage. Further, Prasad et al. (2023) documented a 44.5% increase in sesame yields when SMS was used in conjunction with beneficial microbes. Similarly, Elsakhawy et al. (2020a, b) observed improved agronomic traits and yields in both rice (Oryza sativa) and faba bean (Vicia faba L.). In faba bean, SMS also suppressed Orobanche crenata (broomrape), a parasitic plant that reduces productivity. He et al. (2024) reported that incorporating SMS into soil during rice seedling produc- tion enhanced root development, thereby improving water retention, soil porosity, and nutrient availability. These improvements supported increased P uptake, photosynthesis, and shoot growth. SMS-treated seedlings also ex- hibited higher lateral root numbers, root volume, and surface area, contributing to elevated water-use efficiency. Moreover, Domínguez-Gutiérrez et al. (2022) demonstrated that vermicomposted SMS increases nitrate, K, P, and micronutrient levels, while enhancing cation exchange capacity (CEC). In Hibiscus sabdariffa (Roselle), SMS-amend- ed treatments increased chlorophyll content and biomass, indicating improved nutrient uptake (Ngan and Riddech 2021). A review by Baptista et al. (2023) concluded that the nutrient profile of SMS, especially when co-composted with manure, can match or even surpass that of conventional fertilizers across a wide range of crops. SMS effect on soil SMS application enhances soil physical, chemical, and biological properties, thereby improving overall soil func- tionality and health, although the magnitude of effects varies with treatment, method, and SMS origin. Physical properties Owing to its fibrous organic content, SMS improves soil structure, porosity, aeration, water-holding capacity, and root penetration. These effects, linked to greater aggregation and lower bulk density, also enhance temperature regulation (Marín-Benito et al. 2016, Ma et al. 2025). Nakatsuka et al. (2016) reported the development of granu- lar microstructures and spongy aggregates in SMS-treated soils, providing greater physical stability. Kumar and Chugh (2022) also found improved organic matter content and moisture retention in SMS-amended soils. However, environmental factors can complicate these benefits. For example, in hot, arid climates, SMS may form a surface crust that inhibits germination and moisture penetration (Kahn et al. 2012). Composting and ver- micomposting processes help mitigate these effects by enhancing texture and reducing compaction (Domínguez- Gutiérrez et al. 2022). AGRICULTURAL AND FOOD SCIENCE Agricultural and Food Science (2025) 34: xxx–xxx 5 https://doi.org/10.23986/afsci.163895 Chemical properties SMS contains organic matter but also macro and micronutrients (Mahato et al. 2024, Ma et al. 2025), and SMS has been shown to improve SOC, P, and K availability (Igual et al. 2024). These changes improve nutrient retention and availability (Marín-Benito et al. 2016, Domínguez-Gutiérrez et al. 2022). Although fresh SMS can have high EC and alkaline pH, raising salinity concerns, composting is a proven strategy to stabilize pH, reduce EC, and decrease volatile or phytotoxic compounds. This process also enhances the nutrient profile, reduces potential toxicity, and creates a safer, more consistent organic fertilizer (Medina et al. 2009, Ultra Jr. et al. 2018, Pan et al. 2023, Wang et al. 2024). SMS also enhances buffering capacity, supporting pH balance in the long term (Gupta et al. 2024). Soil fertility, microbial activity, and overall soil health SMS sustains soil fertility by enriching organic matter and releasing nutrients gradually, while stimulating micro- bial biomass, respiration, enzymatic activity, and diversity (Grimm and Wösten 2018, Paula et al. 2020, Martín et al. 2023b, Nie et al. 2024, Ma et al. 2025). Residual fungal mycelia and bioactive compounds further enhance microbial colonization and long-term soil health. Incorporating beneficial organisms such as Trichoderma harzi- anum and Pseudomonas fluorescens amplifies these effects by promoting root development and suppressing soil- borne pathogens (Kumar and Chugh 2022, Prasad et al. 2023, Mahato et al. 2024). Beyond fertility enhancement, SMS contributes to soil remediation by supporting microbial communities that de- grade pollutants and immobilize heavy metals, thereby reducing phytotoxicity and environmental risk (Omoni et al. 2020, Malik et al. 2021, Amarasinghe and Jayaweera 2022, Pan et al. 2023, Gupta et al. 2024). SMS also improves nutrient cycling efficiency by stabilizing organic matter and enhancing the mineralization of N, P, and K (Lou et al. 2017, Baptista et al. 2023). Vermicomposting increases humic substances and nutrient availability, while composting reduces gaseous N losses and boosts overall fertility (Grimm and Wösten 2018, Domínguez-Gutiérrez et al. 2022). Finally, SMS indirectly supports soil fertility by improving plant physiological traits such as chlorophyll content and water retention (Elsakhawy et al. 2020b, Kumar and Chugh 2022) and by promoting beneficial microorganisms— including ligninolytic fungi, actinomycetes, and nutrient-solubilizing rhizobacteria—that enhance soil resilience and nutrient turnover (Ngan and Riddech 2021, Gupta et al. 2024). Long-term agronomic implications SMS has been recognized for its potential as a biofertilizer, organic amendment, and key input in sustainable ag- riculture. Though not all studies provide long-term data, those that do suggest enduring improvements to soil properties and fertility. For instance, Ma et al. (2025) concluded that repeated SMS application improves SOC, mi- crobial activity, and system stability—key indicators of long-term soil health. Lou et al. (2017) reported that SMS releases N gradually, reducing dependency on synthetic inputs. However, Kwiatkowska et al. (2024) suggested that the benefits may diminish if the application is discontinued or not integrated with other strategies. While most studies do not directly measure long-term crop yields, indirect data suggest positive trends. Paula et al. (2020) found that SMS outperformed synthetic fertilizers over 14 weeks, likely due to improved microbial colonization. Some forms of SMS even improve over time as decomposition advances. In addition to its agronom- ic benefits, SMS also facilitates bioremediation. Marín-Benito et al. (2016) and Pan et al. (2023) found that SMS reduces cadmium uptake in crops, improving food safety. Thus, by being reused across mushroom cultivation, composting, animal feed, and fertilization, SMS contributes to the concept of circular agriculture. It closes nutrient loops, reduces waste, and supports sustainable produc- tion systems (Martín et al. 2023a, Mahato et al. 2024, Mwangi et al. 2024). Limitations and challenges While SMS presents numerous agronomic and environmental advantages, several limitations hinder its wide- spread adoption. Key challenges include high EC, variable pH, inconsistent nutrient composition, potential chemical residues, and compatibility issues with certain crops. The key limitations and challenges associated with the SMS application, along with potential mitigation strategies, are summarized in Table 2. AGRICULTURAL AND FOOD SCIENCE Agricultural and Food Science (2025) 34: xxx–xxx 6 https://doi.org/10.23986/afsci.163895 One of the primary concerns is the elevated EC of SMS, due to high concentrations of ions such as K+, Na+, Cl-, and nitrite. High EC levels can induce osmotic stress in plants, inhibit water uptake, and cause nutrient imbalances (Medina et al. 2009, Postinguel et al. 2025). Salinity is particularly detrimental to salt-sensitive crops and young seedlings, which are more vulnerable due to their underdeveloped root systems and limited capacity for water uptake (Lethin et al. 2022, Mousavi et al. 2022). For example, strawberries are susceptible to salt stress, which can damage photosynthetic pigments such as chlorophyll and carotenoids, thereby reducing yield. Additionally, residual organic compounds in SMS may hinder seed germination and initial root growth (Prasad et al. 2021, Velusami et al. 2021, Kwiatkowska and Joniec 2022). Although these issues are significant, mitigation strategies such as leaching or blending SMS with other substrates can help reduce salinity. However, these can raise costs and may not be feasible for all growers (Roy et al. 2015, Paula et al. 2017). Nutrient imbalances also pose a concern. SMS often lacks essential micronutrients, particularly Fe, Zn, and Mn, which are critical for plant growth and enzymatic functions. Iron deficiency, for instance, can limit root elonga- tion and nutrient uptake (Postinguel et al. 2025). Additionally, the high levels of K commonly found in SMS may interfere with Ca uptake, potentially leading to physiological disorders in plants. This emphasizes regular nutrient monitoring and targeted supplementation when required (Zhai et al. 2009, Velusami et al. 2021, Alves et al. 2022). Another important challenge relates to N availability. Fresh SMS often contains a large proportion of organic N bound in recalcitrant lignocellulosic fractions, making it only partially available to plants in the short term (Kahn et al. 2012). Its fertilizer benefit is therefore strongly dependent on composting or vermicomposting processes that mineralize organic N into plant-available forms. Even after such treatments, nitrogen release can be slow and uneven, and some batches with high C:N ratios may still induce temporary N immobilization, reducing short-term availability (Hackett 2015). Additional N losses can occur through ammonia volatilization during storage or surface application, and nitrate leaching after mineralization, particularly in coarse soils, further decreasing N-use efficiency (Wiśniewska-Kadżajan and Malinowska 2022). While composting improves N availability and reduces the risk of immobilization, it requires additional time, labor, and infrastructure, increasing management costs for farmers (Roy et al. 2015). These chal- lenges add to the broader issue of uneven nutrient content and physical structure, making consistent and practi- cal use of SMS as a fertilizer more complex for end-users. Another limitation is the potential presence of residual agrochemicals. SMS may contain traces of pesticides used during mushroom cultivation. These compounds can adversely affect soil microbial communities, pose phytotoxic risks to plants, and even contaminate surrounding soil and water bodies (Postemsky et al. 2016, Paula et al. 2017, Kwiatkowska and Joniec 2022). As a result, conducting soil toxicity assessments is essential to ensure the safe application of SMS in horticultural systems. The physical characteristics of SMS also present challenges. Its properties vary considerably depending on the mushroom species, initial substrate, and composting methods used. This inconsistency makes it difficult to stand- ardize SMS as a commercial growing medium, as batch-to-batch differences can affect plant performance (Prasad et al. 2021, Velusami et al. 2021, Khalil et al. 2024). Moreover, improper storage can cause spoilage, contamination, and compaction, reducing aeration and drainage and complicating the use of SMS for crops with specific needs. However, mixing SMS with porous materials enhances the structure and improves performance (Zied et al. 2020). Moisture retention is another double-edged characteristic. SMS often retains excessive moisture, which can create anaerobic conditions in the root zone. This may encourage harmful microbial activity, promote the growth of fungi and bacteria, and lead to denitrification or waterlogging (Roy et al. 2015, Mousavi et al. 2023). Therefore, maintaining an appropriate balance between drainage and water retention is crucial to preserving soil health and crop productivity (Medina et al. 2009, Zied et al. 2020). Although SMS has demonstrated suppressive effects against several soilborne pathogens, it may also harbor risks. Certain beneficial fungi, such as Trichoderma species, could become opportunistic pathogens in specific horti- cultural settings (Hernández et al. 2021). Incompletely decomposed SMS may release phytotoxic volatile organic compounds, which can negatively affect plant growth (Owaid et al. 2017, He et al. 2024). Moreover, improper disposal of SMS can contribute to GHG emissions—including methane and CO₂—further impacting the environ- ment (Wang et al. 2024). As such, the responsible handling and treatment of SMS are essential for minimizing environmental harm. AGRICULTURAL AND FOOD SCIENCE Agricultural and Food Science (2025) 34: xxx–xxx 7 https://doi.org/10.23986/afsci.163895 Finally, a major obstacle to wider SMS use is the lack of standardized guidelines and regulations. This gap hinders large-scale adoption, especially in commercial horticulture. Developing clear standards and protocols is essential for promoting safe, effective, and region-specific SMS applications (Kousar et al. 2024). SMS as a soil amendment in horticulture Spent mushroom substrate (SMS) has been extensively studied for its potential to enhance plant growth, yield, and quality in horticultural systems. SMS includes key macronutrients that play vital roles in plant development (Medina et al. 2009). Research has demonstrated that incorporating SMS into soil increases microbial activity, SOC, and overall organic matter—factors that are fundamental for healthy plant growth (Hernández et al. 2021). When combined with other organic materials, SMS further improves soil structure and water retention, promot- ing root development and nutrient uptake. These combined effects can lead to significant gains in plant produc- tivity (Oliveira Vieira et al. 2022). Key findings on the use of SMS in horticultural systems—including its effects on nutrient availability, crop productivity, peat replacement, and disease suppression—are summarized in Table 1. SMS effect on horticultural productivity Several studies have demonstrated that SMS can significantly enhance crop productivity, though outcomes de- pend on SMS quality, application method, and environmental conditions. SMS can be considered a biofertilizer in horticultural systems (Paula et al. 2017). Vegetable studies reported yield increases linked to improved root zone conditions and improved soil structure (Martín et al. 2023b). When combined with peat, SMS has also been shown to improve seed germination, early growth, disease resistance, and final seedling weights in a variety of crops, including lettuce (Lactuca sativa), cucumber (Cucumis sativus), tomato (Solanum lycopersicum), and strawberry (Fragaria × ananassa) (Mwangi et al. 2024). SMS has further promoted the growth of sweet pepper (Capsicum annuum), enhancing plant height, branching, and biomass formation (Roy et al 2015). Treatments involving oyster (Pleurotus ostreatus) SMS leachate, weath- ered oyster SMS, and leachate from button mushroom (Agaricus bisporus) compost all yielded better than the control. The best results were seen with the oyster mushroom leachate, followed by weathered oyster SMS and Table 2. Summary of the main limitations and challenges associated with the use of Spent Mushroom Substrate (SMS) in agricultural and horticultural systems, including their potential impacts on crop performance and soil health, along with suggested mitigation strategies and key references Limitation / Challenge Description Potential Mitigation / Notes References High Electrical Conductivity (EC) and variable pH Excess salts (K⁺, Na⁺, Cl⁻, nitrite) can cause osmotic stress, inhibit water uptake, and reduce growth, especially in salt-sensitive crops. Also, an Inconsistent pH can affect nutrient availability and crop performance Leaching, composting, or vermicomposting, blending with other substrates; Adjust pH via blending or liming Medina et al. 2009; Postinguel et al. 2025; Lethin et al. 2022 Nutrient Imbalances Deficiency of micronutrients (Fe, Zn, Mn), high K affecting Ca uptake Targeted supplementation, regular nutrient monitoring Alves et al. 2022; Zhai et al. 2009 Nitrogen Availability Organic N often bound in lignocellulose, slow mineralization; risk of temporary immobilization Composting or vermicomposting, reapplication Kahn et al. 2012; Hackett, 2015; Roy et al. 2015 Residual Agrochemicals Traces of pesticides may affect soil microbes, plants, or water Soil toxicity testing, responsible sourcing, Leaching, composting, or vermicomposting, blending with other substrates Kwiatkowska and Joniec, 2022; Postemsky et al. 2016 Physical Variability Batch-to-batch differences in texture, structure, and porosity; storage issues Blend with porous materials, proper storage Prasad et al. 2021; Zied et al. 2020 Moisture Retention Excess water can create anaerobic conditions, promote harmful microbes Adjust substrate mix and drainage Medina et al. 2009 Potential Phytotoxicity / Opportunistic Pathogens Incompletely decomposed SMS may release toxic compounds; beneficial fungi may act as pathogens Ensure proper composting and monitoring He et al. 2024; Hernández et al. 2021 Environmental Impact Improper disposal may generate GHG emissions Proper treatment and reuse Wang et al. 2024 Lack of Standardization and Guidelines No uniform protocols limiting large-scale adoption Development of standards and regulations Kousar et al. 2024 AGRICULTURAL AND FOOD SCIENCE Agricultural and Food Science (2025) 34: xxx–xxx 8 https://doi.org/10.23986/afsci.163895 button mushroom compost. Consistent with this, another study evaluated SMS as a partial substitute for peat in red baby leaf lettuce production, using three treatments: SMS from A. bisporus (Ab), P. ostreatus (Po), and a 70/30% Ab and Po mixture (AbPo), each mixed with peat at a 1:4 ratio. All SMS treatments outperformed peat alone, with yield increases of up to 2.5-fold under non-infested conditions and up to 7-fold (3 to 7-fold) under Pythium irreg- ulare-infestation. This implies that the remarkable yield response was primarily associated with the suppression of P. irregulare disease pressure. The AbPo mixture showed the most potent effect, reducing disease incidence by 50%, compared to 38% and 15% for Ab and Po alone, respectively (Hernández et al. 2021). These results confirm that SMS can be a sustainable and effective alternative to traditional peat-based substrates. In accordance with these, other studies indicated that applications of SMS compost nearly doubled pineapple (Ananas comosus) fruit yield compared to control treatments in Nigeria (Orluchukwu and Adedokun 2014). Simi- larly, research on root vegetables revealed that radish (Raphanus sativus) yield increased in spring but declined in fall with high SMS application rates—likely due to salt buildup or phytotoxic effects under hot, dry conditions in Oklahoma (Kahn et al. 2012). These results emphasize the importance of optimizing seasonality, compost matu- rity, and dosage. This is further corroborated by Amarasinghe and Jayaweera (2022), who demonstrated that rad- ish germination and root development were improved with N-enriched and microbially inoculated SMS. Further evidence comes from eucalyptus (Eucalyptus spp.) production under water-deficit conditions in Brazil. A study showed that combining SMS with synthetic fertilizers increased shoot weight, collar diameter, and seedling height, while also improving root dry matter and overall plant robustness (Postinguel et al. 2025). These results reinforce the suitability of SMS as a peat substitute, particularly in environments prone to stress. The pathogen-suppressive properties of SMS are attributed to its microbial communities, including endophytes and plant growth-promoting rhizobacteria. These microbes enhance disease resistance by modulating the rhizo- sphere. For instance, SMS has been shown to suppress Fusarium wilt in cucumbers by reshaping root-associated microbial communities (Wang et al. 2020). Similarly, SMS enriched with beneficial microbes improved seedling survival and nutrient uptake in tomato, chili (Capsicum annuum L.), and sesame (Sesamum indicum), by reducing pathogen pressure and enhancing rhizosphere conditions (Kumar and Chugh 2022, Mahato et al. 2024). Addition- ally, SMS can be integrated into co-cultivation systems to optimize resource use. A study demonstrated that to- mato production could be successfully combined with mushroom farming, providing a dual-use system that low- ers production costs while facilitating the benefits of SMS (Oliveira Vieira et al. 2022). Nutrient content and peat replacement Peat is widely used worldwide as a horticultural substrate due to its favorable physical and chemical properties, including high organic carbon content, good aeration, high water-holding capacity, and nutrient retention. More- over, peat’s acidic pH (typically ranging from 3.5 to 4.5) and low electrical conductivity (EC) make it particularly suitable for salt-sensitive crops (Hernández et al. 2021). Peat’s high organic carbon levels contribute to better nutrient retention and improved soil structure (Gao et al. 2015). Peat also contains trace micronutrients such as zinc (Zn), copper (Cu), iron (Fe), and manganese (Mn), which support overall plant health (Postinguel et al. 2025). In contrast, while SMS is lower in organic matter than peat, it is richer in nutrients and exhibits higher microbial activity (Medina et al. 2009). Its pH ranges from slightly acid- ic to alkaline (5.1 to 8.9), and its EC is higher than that of peat. SMS also contains more macronutrients, though typically fewer micronutrients. This variability—due to differences in SMS origin, type, and processing—impacts its performance with salt-sensitive and pH-sensitive crops. However, such variability also enables substrate cus- tomization (Meng et al. 2018, Ravlikovsky et al. 2024). Nonetheless, SMS can sometimes lack essential micronutrients, such as Fe, which are critical for root development and photosynthesis (Kwiatkowska and Joniec 2022). Still, SMS may contain other micronutrients at levels like peat, depending on the feedstock composition, cultivation conditions, processing, and storage techniques (Roy et al. 2015). Hence, using SMS as a peat alternative offers several benefits: SMS provides nutrients comparable to peat with slow-release benefits (Medina et al. 2009, Roy et al. 2015) and improves soil structure, aeration, and water retention. While high pH and EC may stress sensitive crops, leaching can reduce salinity without causing signifi- cant nutrient or microbial losses (Prasad et al. 2021). AGRICULTURAL AND FOOD SCIENCE Agricultural and Food Science (2025) 34: xxx–xxx 9 https://doi.org/10.23986/afsci.163895 Environmental considerations of SMS use The use of SMS offers several environmental benefits, despite some limitations. A comprehensive overview of the environmental benefits, sustainability implications, and limitations of SMS use is presented in Table 1 and 2. Environmental and sustainability implications of SMS use Replacing peat with SMS provides multiple environmental and agronomic benefits by reducing reliance on non- renewable resources, synthetic fertilizers, and agricultural waste. Peatlands, which have accumulated large carbon stocks over millennia, are critical ecosystems for climate regulation and biodiversity. However, peat extraction is energy-intensive, releases stored carbon, and causes habitat destruction, threatening rare species and ecosystem balance (Paula et al. 2017, Meng et al. 2018, Zakaria et al. 2023). SMS, often discarded through dumping, burning, or landfilling, represents a significant opportunity for resource recovery. Mushroom cultivation produces 2.5–5.0 kg of SMS per kg of mushrooms (Sendi et al. 2013, Postinguel et al. 2025). Repurposing this byproduct as a peat alternative diminishes ecological damage from peat mining, lowers GHG emissions, and mitigates environmental risks (Priadi et al. 2018, Kousar et al. 2024, Ravlikovsky et al. 2024). Due to its high organic matter, nutrients, and residual fungal biomass, SMS enriches soils, supports carbon sequestration, enhances microbial activity, and improves overall soil health (Roy et al. 2015, Mohd Hanafi et al. 2018, Othman et al. 2020). It further contributes to water retention, erosion control, remediation of contaminated soils, and resilience of farming systems. Moreover, SMS promotes beneficial rhizosphere microbes, reduces chemi- cal pesticide reliance, and enhances productivity and agroecosystem resilience (Leong et al. 2022). In line with circular economy principles, SMS transforms waste into a valuable input, reducing transport-related emissions when used locally in place of imported peat (Singh et al. 2021, Thygesen et al. 2021, Afsar et al. 2024). Many countries have already restricted peat use for environmental reasons, creating strong demand for viable substitutes such as SMS (Velusami et al. 2021). Altogether, SMS functions as a renewable, multifunctional, and eco-friendly amendment that protects peatlands, improves soil conditions, reduces environmental impacts, and advances climate action, soil restoration, and sustainable farming (Phan and Sabaratnam 2012, Martín et al. 2023a, Ravlikovsky et al. 2024). Carbon sequestration Carbon sequestration is a crucial component of sustainable land management, and SMS plays a significant role in this process. Rich in lignocellulosic polymers—including lignin, cellulose, hemicelluloses—and fungal biomass, SMS decomposes slowly, contributing to long-term accumulation of SOC. As SMS components break down, they are transformed into humic substances through microbial and composting processes, which improve soil structure, nutrient retention, and fertility (Singh et al. 2021, Leong et al. 2022). Composting SMS further stabilizes organic matter, promotes humification, and reduces CO₂ emissions compared to untreated waste. For instance, partially decomposed oyster mushroom SMS increases enzymatic activity, facilitating the breakdown of complex C molecules into stable residues (Afsar et al. 2024). Beyond its chemical composition, SMS promotes microbial growth, thereby influencing soil C dynamics. It boosts microbial biomass and activity, enhances bacterial populations that process organic C, and promotes the formation of soil aggregates. These bioprocesses convert organic inputs into stable forms of soil C, supporting nutrient cycling and long-term carbon storage. As a result, incorporating SMS into soil management systems not only improves soil health but also supports climate resilience by enhancing microbial stabilization, sustaining soil carbon pools, and aligning with broader goals for carbon-neutral farming (Singh et al. 2021, Leong et al. 2022, Khalil et al. 2024). Synthesis of findings As global agriculture shifts toward sustainability, finding alternatives to peat —a carbon-rich, non-renewable ma- terial —has become crucial. Spent mushroom substrate (SMS), a by-product of mushroom cultivation, offers a re- newable substitute that supports circular economy and zero-waste principles. AGRICULTURAL AND FOOD SCIENCE Agricultural and Food Science (2025) 34: xxx–xxx 10 https://doi.org/10.23986/afsci.163895 Benefits of SMS as a soil amendment In farming, SMS improves fertility, soil structure, and microbial activity. Physically, it improves porosity, aeration, and water retention (Nakatsuka et al. 2016, Grimm and Wösten 2018). Chemically, it supplies macro- and micro- nutrients, increases CEC, and buffers pH (Domínguez-Gutiérrez et al. 2022, Martín et al. 2023b). Biologically, it stimulates microbial biomass, enzymatic activity, and respiration, improving nutrient cycling (Paula et al. 2020, Ma et al. 2025), with consistent increases in dehydrogenase and β-glucosidase activity (Kwiatkowska et al. 2024). SMS boosts yield in crops, often matching or outperforming synthetic fertilizers, particularly when combined with manure or compost (Baptista et al. 2023). It also raises organic matter, improves N retention and aggregate sta- bility, and reduces nutrient leaching and heavy metal uptake (Marín-Benito et al. 2016, Pan et al. 2023). Its use reinforces circular economy principles (Martín et al. 2023a, Mwangi et al. 2024). Limitations of SMS as a soil amendment Fresh or poorly composted SMS may contain undecomposed lignocellulose and salts, leading to N immobilization, phytotoxicity, or poor germination (Kahn et al. 2012, Amarasinghe and Jayaweera 2022). High pH and EC levels pose additional risks; leaching, composting, or vermicomposting can stabilize organic matter and reduce salt levels (Ultra Jr. et al. 2018, Domínguez-Gutiérrez et al. 2022). SMS effects are most potent in the first years, especially for microbial activity and nutrient availability, and decline without reapplication or integration into fertility strat- egies (Kwiatkowska et al. 2024). Thus, SMS is best applied within an integrated nutrient management framework. Benefits of SMS as a peat substitute SMS improves plant growth, yield, germination, and soil health by providing high organic matter, beneficial mi- crobes, and a balanced nutrient profile, thereby enhancing soil structure, microbial activity, and root development (Medina et al. 2009, Hernández et al. 2021). SMS also suppresses pathogens through shifts in the rhizosphere mi- crobiome and pathogen competition (Roy et al. 2015, Meng et al. 2018, Hernández et al. 2021). Environmentally, peat extraction drives GHG emissions, wetland loss, and biodiversity decline, while SMS reduces landfill waste and emissions linked to peat mining and fertilizer use (Huang et al. 2022, Zakaria et al. 2023, Postinguel et al. 2025). Limitations of SMS as a peat substitute Fresh SMS often has high EC from soluble salts, stressing salt-sensitive crops like pepper and strawberry; pre- leaching or blending mitigates this but raises costs and labor (Medina et al. 2009, Paula et al. 2017). Nutrient im- balances, particularly micronutrient deficiencies in SMS, require attention (Zhai et al. 2009, Mohd Hanafi et al. 2018). Composition also varies by mushroom species, substrate, and composting, complicating standardization and regulation (Velusami et al. 2021, Wang et al. 2024). The lack of quality protocols and usage guidelines limits the broader adoption of SMS (Medina et al. 2009, Kousar et al. 2024). Knowledge gaps Despite growing interest in using SMS in horticultural and agricultural systems, several knowledge gaps remain to be elucidated to enable its broader adoption. While many short-term studies report benefits, such as improved soil structure, enhanced water retention, increased microbial activity, and higher crop yields, the long-term effects of SMS on soil health, plant performance, and nutrient cycling remain insufficiently explored. Multi-year trials are particularly scarce, making it difficult to assess the sustainability of these benefits across different agroecologi- cal conditions. Some evidence even suggests a decline in positive effects, such as microbial activity and nutrient availability, by the third year without reapplication. This underscores the need for extended field studies to evalu- ate the persistence, variability, and potential risks of SMS use over time. Furthermore, interactions between SMS-derived microbial communities and plant roots have received limited attention. It is unclear whether such interactions could lead to undesirable shifts in the rhizosphere dynamics or long-term impacts on plant health. The variability in SMS composition—driven by differences in mushroom spe- cies, substrates, and storage conditions—also presents challenges. Tailored formulations for specific crop types remain underdeveloped, underscoring the need for targeted research to optimize SMS use across diverse horti- cultural settings. AGRICULTURAL AND FOOD SCIENCE Agricultural and Food Science (2025) 34: xxx–xxx 11 https://doi.org/10.23986/afsci.163895 Concerns also persist regarding the potential for salt accumulation, nutrient imbalances, or the accumulation of trace contaminants, such as heavy metals, particularly when used continuously or repeatedly. Despite growing interest, research into optimal reapplication rates and timing is limited. SMS decomposition and nutrient release rates vary significantly with compost maturity and environmental conditions, such as temperature and moisture. Finally, while SMS holds potential in circular agriculture, its economic feasibility and role in sustainable farming systems remain poorly explored. Practical implications SMS offers several benefits for farming, including improving soil structure and microbial diversity, and reducing nutrient leaching and heavy metal uptake. From a policy standpoint, SMS facilitates the goals of the circular econ- omy and waste reduction. Support mechanisms such as subsidies, certification schemes, and decentralized com- posting programs could encourage its broader use. Commercially, it offers opportunities for mushroom producers and composting companies to generate value-added products, including organic fertilizers and customized growing media. Given its low cost, widespread availability, and environmental benefits, SMS can meaningfully contribute to more resilient and sustainable agri-food systems. However, while SMS is produced in large volumes globally, its application in open-field agriculture faces practical constraints due to the high rates typically required (20–100 t ha⁻¹). Such bulk volumes increase transport, han- dling, and spreading costs, limiting the feasibility of large-scale adoption. To improve scalability, several strategies are worth consideration, including co-composting SMS with other organic wastes to increase nutrient density, pelletizing or drying to reduce bulk and transport costs, and establishing decentralized composting and recycling networks near mushroom farms. These approaches can enhance logistics, reduce environmental footprints, and make SMS a more viable amendment in extensive field systems. Recommendations for future research To realize the full potential of SMS, further research must address key gaps: Standardization and Regulation: The variability of SMS due to differences in mushroom species, cultivation substrates, microbial community found in SMS, and composting processes presents a challenge for consistency. Developing a standardized classification system and regulatory guidelines would support the commercial scaling of this approach. Blending and Formulation: Research should focus on how SMS can be blended with other materials, such as syn- thetic fertilizers or co-composted substrates, to develop flexible and crop-specific formulations that enhance plant performance and nutrient efficiency. Long-Term Trials: Multi-year field studies are crucial for evaluating the long-term persistence of SMS effects across multiple growing seasons, crop rotations, and climatic zones. This includes monitoring for yield sustainability, soil health indicators, and ecological impacts. Risk Assessment and Reapplication Strategies: Further studies are necessary to establish safe and effective reapplica- tion schedules and to monitor potential risks, including salinity, nutrient imbalance, and heavy metal accumulation. Economic and Environmental Evaluation: Life-cycle assessments and techno-economic analyses should be conducted to quantify the costs, benefits, and practical scalability of SMS use across various farming systems, including open-field agriculture, where high application rates currently pose logistical and economic challenges. By addressing these research needs, SMS can transition from an underutilized bioresource into a mainstream in- put for sustainable agriculture and horticulture. Conclusion This review synthesizes current research on agronomic performance and environmental implications of using Spent Mushroom Substrate as a soil amendment and biofertilizer. The findings consistently support SMS as a re- newable and effective alternative to both synthetic fertilizers and non-renewable peat. Rich in organic matter and AGRICULTURAL AND FOOD SCIENCE Agricultural and Food Science (2025) 34: xxx–xxx 12 https://doi.org/10.23986/afsci.163895 nutrients, SMS improves soil structure, porosity, water retention, pH regulation, and nutrient availability, while enhancing microbial activity and disease suppression. It supports nutrient cycling, serves as a carrier for beneficial microbes, and helps degrade agrochemical residues. Compared to peat, SMS is a more sustainable, locally avail- able alternative that supports circular economy principles by reducing waste and conserving natural resources. However, variability in composition, risks of phytotoxicity, and the lack of regulatory standards remain challenges. Future research should emphasize long-term field trials, standardization of processing, and crop-specific guide- lines. Overall, SMS presents a promising strategy for improving soil health, boosting productivity, and advancing sustainable agriculture. With continued research and supportive policy, it can play a central role in regenerative and circular farming systems. References Afsar, M., Zia, A., Us Salam, M.B., Ahmad, M.N., Khan, A.A., ul Haq, T., Aziz, T. & Alasmari, A.F. 2024. A multifaceted analysis of spent mushroom substrate of selected oyster mushrooms for enzymatic activity, proximate composition, and antimicrobial activity. Italian Journal of Food Science 36: 165–174. https://doi.org/10.15586/ijfs.v36i1.2457 Alves, L.d.S., Caitano, C.E.C., Ferrari, S., Vieira Júnior, W.G., Heinrichs, R., de Almeida Moreira, B.R., Pardo-Giménez, A. & Zied, D.C. 2022. Application of Spent Sun Mushroom Substrate in Substitution of Synthetic Fertilizers at Maize Topdressing. Agron- omy 12: 2884. https://doi.org/10.3390/agronomy12112884 Amarasinghe, S.R. & Jayaweera, W.M.C.S. 2022. Evaluation of phytotoxicity of composts produced from spent mushroom substrate using seed germination bioassay of Raphanus sativus L. Tropical Agricultural Research & Extension 25. https://doi.org/10.4038/tare.v25i4.5601 Atallah, E., Zeaiter, J., Ahmad, M.N., Leahy, J.J. & Kwapinski, W. 2021. Hydrothermal carbonization of spent mushroom com- post waste compared against torrefaction and pyrolysis. Fuel Processing Technology 216: 106795. https://doi.org/https://doi. org/10.1016/j.fuproc.2021.106795 Baptista, F., Almeida, M., Paié-Ribeiro, J., Barros, A.N. & Rodrigues, M. 2023. Unlocking the potential of spent mushroom substrate (SMS) for enhanced agricultural sustainability: From environmental benefits to poultry nutrition. Life 13: 1948. https://doi.org/10.3390/life13101948 Courtney, R.G. & Mullen, G.J. 2009. Soil quality and barley growth as influenced by the land application of two compost types. Bioresource Technology 99: 2913–2918. https://doi.org/10.1016/j.biortech.2007.06.034 Domínguez-Gutiérrez, M., Gaitán-Hernández, R., Moctezuma-Pérez, I., Barois, I. & Domínguez, J. 2022. Stabilization of spent mush- room substrate by composting and vermicomposting: Nutrient dynamics and suitability for soil amendment. Emirates Journal of Food and Agriculture 33: 743–753. https://doi.org/10.9755/ejfa.2022.v34.i3.2828 Elsakhawy, T., ALKahtani, M.D.F., Sharshar, A.A.H., Attia, K.A., Hafez, Y.M. & Abdelaal, K.A.A. 2020a. Efficacy of Mushroom Me- tabolites (Pleurotus ostreatus) as A Natural Product for the Suppression of Broomrape Growth (Orobanche crenata Forsk) in Faba Bean Plants. Plants 9: 1265. https://doi.org/10.3390/plants9101265 Elsakhawy, T. & Abd El-Rahem, W. 2020b. Evaluation of Spent Mushroom Substrate Extract as a Biofertilizer for Growth Improve- ment of Rice (Oryza sativa L.). Egyptian Journal of Soil Science 60: 31–42. https://doi.org/10.21608/ejss.2019.18835.1320 Gao, W., Liang, J., Pizzul, L., Feng, X.M., Zhang, K. & Castillio, M.D.P. 2015. Evaluation of spent mushroom substrate as substitute of peat in Chinese biobeds. International Biodeterioration & Biodegradation 98: 107–112. https://doi.org/10.1016/j.ibiod.2014.12.008 Grimm, D. & Wösten, H.A.B 2018. Mushroom cultivation in the circular economy. Applied Microbiology and Biotechnology 102: 7795–7803. https://doi.org/10.1007/s00253-018-9226-8 Gupta, G., Maurya, S., Jha, P.N. & Chauhan, P.S. 2024. Valorization of Spent Mushroom Substrate (SMS) as a sustainable approach to remediation of xenobiotic compounds in groundwater –A comprehensive review. Groundwater for Sustainable Development 26: 101290. https:/doi.org/10.1016/j.gsd.2024.101290 Hackett, R. 2015. Spent mushroom compost as a nitrogen source for spring barley. Nutrient Cycling in Agroecosystems 102: 253– 263. https://doi.org/10.1007/s10705-015-9696-3 He, J., Zheng, G., Liu, Z., Guo, Z., Zhang, W., Li, Y., Zhou, Y. & Xu, H. 2024. Replacing traditional nursery soil with spent mushroom substrate improves rice seedling quality and soil substrate properties. Environmental Science and Pollution Research 31: 39625– 39636. https://dx.doi.org/10.1007/s11356-024-33723-x Hernández, D., Ros, M., Carmona, F., Saez-Tovar, J.A & Pascual, J.A 2021. Composting Spent Mushroom Substrate from Agaricus bisporus and Pleurotus ostreatus Production as a Growing Media Component for Baby Leaf Lettuce Cultivation under Pythium ir- regulare Biotic Stress. Horticulturae 7: 13. https://doi.org/10.3390/horticulturae7020013 Huang, Z., Guan, H., Zheng, H., Wang, M., Xu, P., Dong, S., Yang, Y. & Xiao, J. 2022. Novel liquid organic fertilizer: A potential way to effectively recycle spent mushroom substrate. Journal of Cleaner Production 376: 134368. https://doi.org/10.1016/j. jclepro.2022.134368 Igual, J.M., Andrades, M.S., Frade, C., Carpio, M.J., Marín-Benito, J.M., Rodríguez-Cruz, M.S. & Valverde, A. 2024. Impacts of spent mushroom substrate amendment and season on microbial communities in a semiarid vineyard soil. Applied Soil Ecology 203: 105689. https:/doi.org/10.1016/j.apsoil.2024.105689 Kahn, B.A., Payton, M.E. & Graetz, D.A. 2012. Compost Treatments Interact with Other Factors To Affect Red Radish Production. Compost Science & Utilization 20: 79–86. https://doi.org/10.1080/1065657X.2012.10737029 AGRICULTURAL AND FOOD SCIENCE Agricultural and Food Science (2025) 34: xxx–xxx 13 https://doi.org/10.23986/afsci.163895 Khalil, S., Panda, P., Ghadamgahi, F., Barreiro, A., Rosberg, A.K., Karlsson, M. & Vetukuri, R.R. 2024. Microbial potential of spent mushroom compost and oyster substrate in horticulture: Diversity, function, and sustainable plant growth solutions. Journal of Environmental Management 357: 120654. https://doi.org/10.1016/j.jenvman.2024.120654 Kousar, A., Khan, H.A., Farid, S., Zhao, Q.B. & Zeb, I. 2024. Recent advances on environmentally sustainable valorization of spent mushroom substrate: A review. Biofuels Bioproducts & Biorefining-Biofpr 18: 639–651. https://doi.org/10.1002/bbb.2559 Klausen, S.J., Romero-Soto, L.A., Liaqat, A., Dehghanmanshadi, Z., Strætkvern, K.O., Xiong, S. & Martín, C. 2025. Biorefining spent substrates of shiitake (Lentinula edodes) and oyster mushroom (Pleurotus ostreatus): enzymatic saccharification of cellulose and xylan, with lignin recovery from residues. Bioengineered 16: 2536443. https://doi.org/10.1080/21655979.2025.2536443 Kumar, S. & Chugh, R.K. 2022. Impact of Spent Mushroom Substrate Enriched with Trichoderma harzianum on Damping off Dis- ease in Chilli and Tomato. International Journal of Economic Plants 9: 264–270. https://doi.org/10.23910/2/2022.0470a Kwiatkowska, E. & Joniec, J. 2022. Effects of Agricultural Management of Spent Mushroom Waste on Phytotoxicity and Microbio- logical Transformations of C, P, and S in Soil and Their Consequences for the Greenhouse Effect. International Journal of Environ- mental Research and Public Health 19: 12915. https://doi.org/10.3390/ijerph191912915 Kwiatkowska, E., Joniec, J., Kwiatkowski, C.A., Kowalczyk, K., Nowak, M. & Leśniowska-Nowak, J. 2024. Assessment of the impact of spent mushroom substrate on biodiversity and activity of soil bacterial and fungal populations based on classical and modern soil condition indicators. International Agrophysics 38: 139–154. https://doi.org/10.31545/intagr/184175 Leong, Y.K., Ma, T.W., Chang, J.S. & Yang, F.C. 2022. Recent advances and future directions on the valorization of spent mushroom substrate (SMS): A review. Bioresource technology 344(Pt A): 126157. https://doi.org/10.1016/j.biortech.2021.126157 Lethin, J., Byrt, C., Berger, B., Brien, C., Jewell, N., Roy, S., Mousavi, H., Sukumaran, S., Olsson, O. & Aronsson, H. 2022. Improved Salinity Tolerance-Associated Variables Observed in EMS Mutagenized Wheat Lines. International Journal of Molecular Sciences 23: 11386. https://doi.org/10.3390/ijms231911386 Lou, Z., Sun, Y., Zhou, X., Baig, S.A., Hu, B. & Xu, X. 2017. Composition variability of spent mushroom substrates during con- tinuous cultivation, composting process and their effects on mineral nitrogen transformation in soil. Geoderma 307: 30–37. https://doi.org/10.1016/j.geoderma.2017.07.033 Ma, X., Yan, S. & Wang, M. 2025. Spent mushroom substrate: A review on present and future of green applications. Journal of Environmental Management 373: 123970. https://doi.org/10.1016/j.jenvman.2024.123970 Mahato, D., Khamari, B. & Sahoo, J.P. 2024. Valorization of Spent Oyster Mushroom Substrate with Trichoderma asperellum for Suppression of Sesame Root Rot In Vitro. Waste Biomass Valor 15: 755–769. https://doi.org/10.1007/s12649-023-02190-9 Malik, N.A., Kumar, J., Wani, M.S., Tantray, Y.R. & Ahmad, T. 2021. Role of Mushrooms in the Bioremediation of Soil. In: Dar, G.H., Bhat, R.A., Mehmood, M.A. & Hakeem, K.R. (eds.). Microbiota and Biofertilizers 2: 77–102. https://doi.org/10.1007/978-3-030-61010-4_4 Marín-Benito, J.M., Sánchez-Martín, M.J. & Rodríguez-Cruz, M.S. 2016. Impact of Spent Mushroom Substrates on the Fate of Pesticides in Soil, and Their Use for Preventing and/or Controlling Soil and Water Contamination: A Review. Toxics 4: 17. https://doi.org/10.3390/toxics4030017 Martín, C., Xiong, S., Passoth, V., Põldmaa, K., Solberg, S.Ø., Hultberg, M., Strætkvern, K.O., Golovko, O., Pilotto, F., Müller, B., Pent, M., Klausen, S.J. & Romero-Soto, L.A. 2023a. Mushrooms for enhanced agriculture sustainability – the MUSA concept. C3- BIOECONOMY: Circular and Sustainable Bioeconomy 4: 131–147. https://doi.org/10.21071/c3b.vi4.16591 Martín, C., Zervakis, G.I., Xiong, S., Koutrotsios, G. & Strætkvern, K.O. 2023b. Spent substrate from mushroom cultivation: ex- ploitation potential toward various applications and value-added products. Bioengineered 14. https://doi.org/10.1080/216559 79.2023.2252138 Martín, C., Zervakis, G.I., Xiong, S., Koutrotsios, G. & Strætkvern, K.O. 2023b. Spent substrate from mushroom cultivation: ex- ploitation potential toward various applications and value-added products. Bioengineered 14: 2252138. https://doi.org/10.1080/216559 79.2023.2252138 Meng, X., Dai, J., Zhang, Y., Wang, X., Zhu, W., Yuan, X., Yuan, H. & Cui, Z. 2018. Composted biogas residue and spent mush- room substrate as a growth medium for tomato and pepper seedlings. Journal of Environmental Management 216: 62–69. https://doi.org/10.1016/j.jenvman.2017.09.056 Meshram, V., Mahish, P.K. & Chandrawanashi, N.K. 2024. Mushroom cultivation and application in green technology. In: Microbial Approaches for Sustainable Green Technologies, CRC Press. https://doi.org/10.1201/9781003407683-16 Mohd Hanafi, F.H., Rezania, S., Mat Taib, S., Md Din, M., Yamauchi, M., Sakamoto, M., Hara, H., Park, J. & Ebrahimi, S.S. 2018. En- vironmentally sustainable applications of agro-based spent mushroom substrate (SMS): an overview. Journal of Material Cycles and Waste Management 20: 1383–1396. https://doi.org/10.1007/s10163-018-0739-0 Mousavi, H., Cottis, T. & Solberg, S.Ø. 2023. Nitrogen Enriched Organic fertilizer (NEO) elevates nitrification rates short- ly after application but has no lasting effect on nitrification in agricultural soils. Agricultural and Food Science 32: 179–194. https://doi.org/10.23986/afsci.131722 Mousavi, H., Bruning, B., Van Straten, G., Almås, Å.R., Lethin, J., Naik, N., Hassan, S., Olsson, O. & Aronsson, H. 2022. Effects of In- creasing Salinity by Drip Irrigation on Total Grain Weight Show High Yield Potential of Putative Salt-Tolerant Mutagenized Wheat Lines. Sustainability 14: 5061. https://doi.org/10.3390/su14095061 Mwangi, W.W., Mustafa, M., Kappel, N., Csambalik, L. & Szabó, A. 2024. Practical applications of spent mushroom compost in cultivation and disease control of selected vegetable species. Journal of Material Cycles and Waste Management 26: 1918–1933. https://doi.org/10.1007/s10163-024-01969-9 Nakatsuka, H., Oda, M., Hayashi, Y. & Tamura, K. 2016. Effects of fresh spent mushroom substrate of Pleurotus ostreatus on soil micromorphology in Brazil. Geoderma 269: 54–60. http://dx.doi.org/10.1016/j.geoderma.2016.01.023 AGRICULTURAL AND FOOD SCIENCE Agricultural and Food Science (2025) 34: xxx–xxx 14 https://doi.org/10.23986/afsci.163895 Ngan, N.M. & Riddech, N. 2021. 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 and Biomass Valorization 12: 3801–3811. https://doi.org/10.1007/s12649-020-01278-w Nie, H., Li, C., Jia, Z., Cheng, X., Liu, X., Liu, Q., Chen, M., Ding, Y. & Zhang, J. 2024. Microbial inoculants using spent mushroom substrates as carriers improve soil multifunctionality and plant growth by changing soil microbial community structure. Journal of Environmental Management 370: 122726. https:/doi.org/10.1016/j.jenvman.2024.122726 Oliveira Vieira, V., Almeida Conceição, A., Raisa Barbosa Cunha, J., Enis Virginio Machado, A., Gonzaga de Almeida, E., Souza Dias, E., Magalhães, A.L., Miller, R. & Goncalves, D.S.F. 2022. A new circular economy approach for integrated production of tomatoes and mushrooms. Saudi Journal of Biological Sciences 29: 2756–2765. https://doi.org/10.1016/j.sjbs.2021.12.058 Omoni, V.T., Lag-Brotons, A.J. & Semple, K.T. 2020. Impact of organic amendments on the development of 14C-phenanthrene catabolism in soil. International Biodeterioration & Biodegradation 151: 104991. https://doi.org/10.1016/j.ibiod.2020.104991 Orluchukwu, J.A. & Adedokun, O.M. 2014. Comparative effects of poultry manure and spent mushroom substrate on the growth and yield of pineapple (Ananas comosus) in Nigeria. African Journal of Agricultural Research 9: 2041–2044. https://doi.org/10.5897/AJAR2013.8079 Othman, N.Z., Sarjuni, M.N.H., Rosli, M.A., Nadri, M.H., Yeng, L.H., Ying, O.P. & Sarmidi, M.R 2020. Spent Mushroom Substrate as Biofertilizer for Agriculture Application. In: Zakaria, Z., Boopathy, R. & Dib, J. (eds). Valorisation of Agro-industrial Residues – Vol- ume I: Biological Approaches. Cham: Springer International Publishing. https://doi.org/10.1007/978-3-030-39137-9_2 Owaid, M.N., Abed, I.A. & Al-Saeedi, S.S.S. 2017. Applicable properties of the bio-fertilizer spent mushroom substrate in organ- ic systems as a byproduct from the cultivation of Pleurotus spp. Information Processing in Agriculture 4: 78–82. https://doi.org/10.1016/j.inpa.2017.01.001 Pan, C.X., Deng, T.F., Zhang, L., Ge, L.J., Li, L.Q., Yang, L.S. & Yang, X.S. 2023. Epimedium Herbal Residue as a Bulking Agent for Lignite and Spent Mushroom Substrate Co-composting. Waste and Biomass Valorization 14: 2547–2555. https://doi.org/10.1007/s12649-022-02018-y Paula, F.S., Tatti, E., Abram, F., Wilson, J. & O’Flaherty, V. 2020. Soil prokaryotic community resilience, fungal colonisation and in- creased cross-domain co-occurrence in response to a plant-growth enhancing organic amendment. Soil Biology and Biochemistry 149: 107937. https://doi.org/10.1016/j.soilbio.2020.107937 Paula, F.S., Tatti, E., Abram, F., Wilson, J. & O’Flaherty, V. 2017. Stabilisation of spent mushroom substrate for application as a plant growth-promoting organic amendment. Journal of Environmental Management 196: 476–486. https://doi.org/10.1016/j.jenvman.2017.03.038 Phan, C.W. & Sabaratnam, V. 2012. Potential uses of spent mushroom substrate and its associated lignocellulosic enzymes. Ap- plied Microbiology and Biotechnology 96: 863–873. https://doi.org/10.1007/s00253-012-4446-9 Postemsky, P.D., Marinangeli, P.A. & Curvetto, N.R. 2016. Recycling of residual substrate from Ganoderma lucidum mushroom cultivation as biodegradable containers for horticultural seedlings. Scientia Horticulturae 201: 329–337. https://doi.org/10.1016/j.scienta.2016.02.021 Postinguel, R.D.B., Alves, L.D.S., Ré, B.B., Caitano, C.E.C., Barretto, V.C.D.M. & Zied, D.C. 2025. Spent mushroom substrate for euca- lyptus seedling growth under varying water deficiency conditions. New Forests 56. https://doi.org/10.1007/s11056-024-10082-6 Prasad, D., Singh, S. & Upadhyay, U. 2023. Integrated Approaches for Management of Phytophthora Blight in Sesame (Sesamum indicum L.). International Journal of Economic Plants 10: 143–148. https://doi.org/10.23910/2/2023.0513d Prasad, R., Lisiecka, J., Antala, M. & Rastogi, A. 2021. Influence of Different Spent Mushroom Substrates on Yield, Morphologi- cal and Photosynthetic Parameters of Strawberry (Fragaria × ananassa Duch.). Agronomy 11: 2086. https://doi.org/10.3390/agronomy11102086 Priadi, D. & Saskiawan, I. 2018. The utilization of spent oyster mushroom substrates into compost and its effect on the growth of Indian mustard (Brassica juncea (L.) Czern.) in the screenhouse. International Journal of Agricultural Technology 14: 351–362. https://www.scopus.com/inward/record.uri?eid=2-s2.0-85063542786&partnerID=40&md5=e021ce08af2a79c39d6e7cdb3c88 235a Ravlikovsky, A., Pinheiro, M.N.C., Dinca, L., Crisan, V. & Symochko, L. 2024. Valorization of Spent Mushroom Substrate: Establish- ing the Foundation for Waste-Free Production. Recycling 9: 13. https://doi.org/10.3390/recycling9030044 Roy, S.B., Barman, S., Chakraborty, U. & Chakraborty, B. 2015. Evaluation of Spent Mushroom Substrate as biofertilizer for growth improvement of Capsicum annuum L. Journal of Applied Biology & Biotechnology 3: 22–27. https://doi.org/10.7324/JABB.2015.3305 Sendi, H., Mohamed, M.T., Anwar, M.P. & Saud, H.M 2013. Spent mushroom waste as a media replacement for peat moss in Kai- Lan (Brassica oleracea var. Alboglabra) production. Scientific World Journal 2013: 258562. https://doi.org/10.1155/2013/258562 Singh, G., Tiwari, A., Rathore, H., Prasad, S., Hariprasad, P. & Sharma, S. 2021. Valorization of Paddy Straw Using De-oiled Cakes for P. ostreatus Cultivation and Utilization of Spent Mushroom Substrate for Biopesticide Development. Waste and Biomass Val- orization 12: 333–346. https://doi.org/10.1007/s12649-020-00957-y Thygesen, A., Tsapekos, P., Alvarado-Morales, M. & Angelidaki, I. 2021. Valorization of municipal organic waste into purified lactic acid. Bioresource Technology 342: 125933. https:/doi.org/10.1016/j.biortech.2021.125933 Ultra Jr, V.U., Ong Sotto, J.M.E. & Punzalan, M.R. 2018. Nutrient supplying potential of different spent mushroom substrate preparations in potting media. IOP Conf. Series: Earth and Environmental Science 120: 012010. https://doi.org/10.1088/1755-1315/120/1/012010 Velusami, B., Jordan, S.N., Currant, T. & Grogan, H. 2021. Fertiliser characteristics of stored spent mushroom substrate as a sus- tainable source of nutrients and organic matter for tillage, grassland and agricultural soils. Irish Journal of Agricultural and Food Research 60: 1–11. https://doi.org/10.15212/ijafr-2020-0121 AGRICULTURAL AND FOOD SCIENCE Agricultural and Food Science (2025) 34: xxx–xxx 15 https://doi.org/10.23986/afsci.163895 Wang, S., DU, X., Yin, R., Sun, H., Song, B., Han, Q., Wang, J. & Huang, Y. 2024. Performance of co-composting Pholiota nameko spent mushroom substrate and pig manure at different proportions: Chemical properties and humification process. Journal of Environmental Management 372. https://doi.org/10.1016/j.jenvman.2024.123325 Wang, H.-W., Xu, M., Cai, X.-Y., Feng, T. & Xu, W.-L. 2020. Application of spent mushroom substrate suppresses Fusarium wilt in cucumber and alters the composition of the microbial community of the cucumber rhizosphere. European Journal of Soil Biology 101: 103245. https://doi.org/10.1016/j.ejsobi.2020.103245 Wiśniewska-Kadżajan, B. & Malinowska, E. 2022. The Effects of Spent Mushroom Substrate on the Yield and Nutritional Value of Festulolium braunii (K. Richt.) A. Camus. Agriculture 12: 1537. https://doi.org/10.3390/agriculture12101537 Yagüe, M.R. & Lobo, M.C. 2024. Valorization of organic residues for lettuce production minimizes nitrogen loss compared to min- eral fertilization. Journal of Plant Nutrition 47: 1680–1696. https://doi.org/10.1080/01904167.2024.2319161 Zakaria, Z., Seng, T.T.C., Zailani, S.N., Zabidi, K.A.A. & Salim, S.K. 2023. Utilization of Spent Mushroom Compost in Grey Oyster Mush- room Cultivation. In: Shukor, H., Halim, H.N.A., Ong, H.L., Lee, B.B. & Pisal, M.H.M. (eds). Emerging Technologies for Future Sus- tainability. ICoBiomasSE 2022. Green Energy and Technology. Springer, Singapore. https://doi.org/10.1007/978-981-99-1695-5_23 Zhai, Z., Ehret, D.L., Forge, T., Helmer, T., Lin, W., Dorais, M. & Papadopoulos, A.P. 2009. Organic Fertilizers for Greenhouse Pro- ductivity and Substrate Microbiology. Horticultural Science 44: 800–809. https://doi.org/10.21273/hortsci.44.3.800 Zhu, H., Zhao, S., Yang, J., Meng, L., Luo, Y., Hong, B., Cui, W., Wang, M. & Liu, W. 2018. Growth, Nutrient Uptake, and Foliar Gas Exchange in Pepper Cultured with Un-composted Fresh Spent Mushroom Residue. Notulae Botanicae Horti Agrobotanici Cluj- Napoca 47: 227–236. https://doi.org/10.15835/nbha47111307 Zied, D.C., Sánchez, J.e., Noble, R. & Pardo-Giménez, A. 2020. Use of Spent Mushroom Substrate in New Mushroom Crops to Pro- mote the Transition towards A Circular Economy. Agronomy 10: 1239. https://doi.org/10.3390/agronomy10091239 AGRICULTURAL AND FOOD SCIENCE Agricultural and Food Science (2025) 34: xxx–xxx 16 https://doi.org/10.23986/afsci.163895 Appendix 1 Table 1. Complete final search including search terms, filters, and the number of hits from the database Scopus. Database: Scopus Search Term Limitations (filters, limits, refine) Amount of Hits # 1 TOPIC: (“Spent Mushroom Substrate”) Refined by: Languages (ENGLISH) and publication years: (AFTER 2009) 55 620 # 2 TOPIC: (“Spent Mushroom Substrate as biofertilizer” OR “Spent Mushroom Substrate for soil amendment”) Refined by: Languages (ENGLISH) and publication years: (AFTER 2009) 6 # 3 TOPIC: (“Utilisation of SMS” OR “SMS in agriculture” OR “SMS in horticulture”) Refined by: Languages (ENGLISH) and publication years: (AFTER 2009) 190 # 4 TOPIC: (“Spent Mushroom Substrate” OR “Biofertilizer” OR “Soil Amendment”) Refined by: Languages (ENGLISH) and publication years: (AFTER 2009) 33 774 # 5 TOPIC: (“SMS in agriculture” OR “SMS in horticulture”) Refined by: Languages (ENGLISH) and publication years: (AFTER 2009) 28 # 6 TOPIC: (“Spent Mushroom Substrate” OR “Peat Substitute” OR “Peat”) Refined by: Languages (ENGLISH) and publication years: (AFTER 2009) 133 061 # 7 TOPIC: (“Spent Mushroom Substrate” OR “Growing Media”) Refined by: Languages (ENGLISH) and publication years: (AFTER 2009) 9 451 # 8 TOPIC: (“Spent Mushroom Substrate” OR “SMS and Environment”) Refined by: Languages (ENGLISH) and publication years: (AFTER 2009) 980 Table 2. Complete final search including search terms, filters, and the number of hits from the database Web of Science. Database: Web of Science Search Term Limitations (filters, limits, refine) Amount of Hits # 1 TOPIC: (“Spent Mushroom Substrate”) Refined by: Languages (ENGLISH) and publication years: (AFTER 2009) 17 280 # 2 TOPIC: (“Spent Mushroom Substrate” AND “Biofertilizer*” OR “Soil amendment” OR Soil*) Refined by: Languages (ENGLISH) and publication years: (AFTER 2009) 760 929 # 3 TOPIC: (“Spent Mushroom Substrate” AND “Utilization in agriculture” OR “Utilization in horticulture”) Refined by: Languages (ENGLISH) and publication years: (AFTER 2009) 814 # 4 TOPIC: (“Spent Mushroom Substrate” AND “Biofertilizer” OR “Soil Amendment”) Refined by: Languages (ENGLISH) and publication years: (AFTER 2009) 10 349 # 5 TOPIC: (“Spent Mushroom Substrate” AND Agriculture* OR Horticulture*) Refined by: Languages (ENGLISH) and publication years: (AFTER 2009) 255 977 # 6 TOPIC: (“Spent Mushroom Substrate” AND “Peat Substitute” OR “Peat”) Refined by: Languages (ENGLISH) and publication years: (AFTER 2009) 21 000 # 7 TOPIC: (“Spent Mushroom Substrate” AND “Growing Media”) Refined by: Languages (ENGLISH) and publication years: (AFTER 2009) 2 727 # 8 TOPIC: (“Spent Mushroom Substrate” AND Environment”) Refined by: Languages (ENGLISH) and publication years: (AFTER 2009) 715