2024.01.02.15

Edible mushroom production in Ecuador: opportunities for biotechnological use of agricultural by-products

Laura Scalvenzi¹, Mayra Vanessa Chiriboga Ruilova¹, Estela Guardado Yordi¹, Matteo Radice ¹,

Renier Abreu Naranjo ¹, Amaury Pérez Maertínez¹*

¹Universidad Estatal Amazónica / Puyo / Ecuador.

* Correspondence: amperez@uea.edu.ec; Tel.: 00 593 995629543

Available from. http://dx.doi.org/10.21931/BJ/2024.02.01.15

ABSTRACT

Recent studies estimate that there are between 1.5 and 5.1 million fungi species on Earth, and 150.000 of them have been taxonomically classified. Approximately 2.000 are suitable for food and medicinal applications, but only 35 are cultivated commercially. In Ecuador, 100.000 species are available, although only 5.000 have been taxonomically characterized. Mycoculture is the production of fungi under controlled conditions. Edible mushrooms are saprophytes, and they grow on decomposing organic substrates, mainly on lignocellulosic agricultural by-products such as sawdust, straw, bran and bagasse. Mushroom cultivation turns waste into resource, reducing pollution. Ecuador is an agriculture-based economy, and many lignocellulosic wastes are produced from permanent crops such as cocoa, African palm, sugar cane and coconut. Ecuador's inhabitants are looking for healthy and environmentally friendly products, leading to an increase in demand for locally produced mushrooms, which show high nutritional values and medicinal properties. At the global level, current studies show that edible mushroom production moves around 42 billion dollars annually, projected to 62 billion by 2023. China is the leading producer, with more than 7 million tons per year reported in 2016, followed by the United States, Italy, France and Spain. The increasing rate of edible mushroom global production stands at about 11%.

Keywords: fungi; waste; transformation; biotechnology; mycoprotein

INTRODUCTION

The Earth is home to millions of fungi species (1.5 to 5.1), most still unknown. Considering these estimations, just 2.9% to 10% have undergone taxonomic classification (150.000) ¹. Of known species, 1.3% (2.000) are used daily by people worldwide for food and medicinal purposes, but just 1.7% (35) are intentionally cultivated for commercial reasons ². Researchers estimate that around 100.000 species were found in Ecuador, even though only 5.000 have been taxonomically characterized ³. In tropical America, edible mushroom cultivation started in Mexico in 1933, followed in the forties by Argentina, in the fifties by Colombia, Brazil and Chile, in the sixties by Guatemala, Peru, Ecuador, Venezuela and Costa Rica and the eighties by Bolivia ⁴. Cultivated edible fungi are saprophytes; they grow on decomposing organic substrates, mainly lignocellulosic agricultural by-products such as sawdust, straw, bran, and bagasse. Mushroom cultivation turns waste into resource, reducing pollution caused by them. At the global level, approximately 998 million agricultural waste is produced each year due to production intensification. That increase causes improper waste management and pollution, affecting people and the environment ⁵. The production of edible mushrooms under controlled conditions is known as my-coculture, an affordable alternative to harvest from the natural environment. High volumes of fresh mushrooms are achieved through accessible technologies, allowing easier consumption in the diet, especially for those living in rural areas ⁶. World production of edible mushrooms has increased from 1 to 34 million tons per year from 1978 - 2013. Current studies show that this production moves around 42 billion dollars annually, projected to reach 62 billion dollars by 2023 ⁷. China stands out as the leading producer of edible mushrooms, with more than 7 million tons per year reported in 2016, followed by the United States, Italy, France and Spain⁸. On the other hand, the increasing rate of global production of edible mushrooms stands at about 11% ⁷.

NUTRITIONAL VALUE OF EDIBLE MUSHROOM

Mushrooms are consumed worldwide for their nutritional value. Over 200 species are used as food, although 35 are cultivated commercially. The most cultivated mushrooms, which produce edible carpophores, belong to the Phylum Basidiomycota. The most cultivated genera in the world are Agaricus, Flammulina, Lentinula and Pleurotus ². Mushrooms are rich in water (80-95%), low in calories (2-3% lipid dry weight) and high in proteins (20-25% dry weight) (Table 1). They are a source of minerals such as potassium, phosphorus, magnesium, calcium, copper and zinc, as well as carbohydrates such as chitin, glycogen and hemicelluloses ². Edible mushroom production is environmentally, technologically, and socially sustainable since solid-state fermentation is used on agricultural wastes ⁸. In addition to their high nutritional value, edible mushrooms have also medicinal properties. Bioactive compounds are found in fruiting bodies and mycelium of some species ⁹. Recent studies in India showed the presence of gallic acid, hydroxybenzoic acid, quercetin and epicatechin in some species; those compounds have shown beneficial biological activities for disease prevention ¹⁰. Edible mushrooms have also been investigated for the presence of bioactive compounds such as peptides, polyphenolic compounds, polysaccharides, oligosaccharides, dietary fibers, amino acids and micronutrients, which seems to be able to explain their biological effects ¹¹, such as anti-inflammatory properties ¹², depressive disorder ¹³, anti-obesity treatment ¹⁴ and diabetes mellitus ¹⁵. Due to those effects, they have potential as nutraceuticals ^16,17. Considering the beneficial impacts on human health, edible mushrooms are also considered functional foods ¹⁸. As reported in some research ^19-21, promising in vitro and in vivo studies revealed anti-tumoral, neuroprotective, hepatoprotective, cardioprotective, immune-potentiating and hypocholesterolemic activities. A specific focus about the pivotal role of polysaccharides on gut microbiota regulation have been mentioned in two investigations ^22,23, addressing the prebiotic activity of mushrooms on that specific microbiota, which enhance health benefits for the host. Also, studies on b-glucans suggested a potential clinical use of edible mushrooms in recurrent infections, cardiovascular and metabolic diseases ^24,25. Edible mushrooms have been proposed also as a source of nature-based nutraceutical compounds, which seem to be able to mitigate neurodegenerative diseases by boosting the immune-modulating response. These studies open a new trend research to ameliorate the life of patients with dementia and Alzheimer’s disease and to develop new food supplements or diet strategies for brain and nervous system health ^26,27. However, consumption safety must always be guaranteed, and they must be microbiologically safe and free of toxic compounds. Production techniques must allow the conservation of mushroom nutritional and health benefit value

^28,29.

Mushrooms species: A.b (Agaricus bisporus), A.bl (Agaricus blazei), A.su (Agaricus subrufescens), A.sy (Agaricus sylvaticus), F.v (Flammulina velutipes), L.e (Lentinula edodes), P.e (Pleurotus eryngii), P.d (Pleurotus djamor), P.o (Pleurotus ostreatus). S: dry g 100^-1 | Ss: dry % | F: fresh % nd: not available

Table 1. Nutritional value of some edible mushrooms cultivated worldwide.

MUSHROOMS CULTIVATED IN ECUADOR

Native people from the coast and the Andean and Amazonian regions of Ecuador are used to ingesting mushrooms that grow in the wild. They have developed a traditional knowledge of ethnomycology, including myths as a cultural result of fungi consumption. Gamboa-Trujillo carried out studies on mushroom consumption in Ecuadorian ethnic groups distributed all around the country. Among them, eleven groups consume edible fungi of about 37 species belonging to Basidio- and Ascomycota; all have vernacular names. The most common genera were Agaricus, Auricularia, Coprinus, Favolus, Gymnopus, Lentinus, Phillipsia, Pleurotus, Poliporus, Trametes, Volvariella ^{41, 42}. Despite the vast mushroom consumption by Native people and the high fungus biodiversity, Ecuador is slowly developing large-scale mushroom production for retail trade. The production of the mass market began in the sixties of the past century. AMCESA and KENNET cultivated A. bisporus mushroom, the most common edible Mushroom in the Ecuadorian market ³.

Today, mushroom production and industrialization are mainly centered in Pichincha province (Figure 1). Ecuadorian consumer preferences have significantly influenced the international sales of the country's mushroom production. In 1968, Ecuador began cultivating mushrooms. Of the first harvest, which totaled about 45 tons, roughly 90% were exported, while the remainder was sold locally ⁴³. In Ecuador, the most cultivated species are the oyster mushroom (P. ostreatus), pink oyster mushroom (P. djamor), white Mushroom (A. bisporus) and shiitake (L. edodes). Most of the production is carried out by privates or through rural development projects sponsored by the government ⁴⁴. To our knowledge, little data is available about Ecuador's current edible mushroom market; the limited information is about dried mushrooms. Data revealed a clear growth trend in Ecuador's dried mushroom production from 2007 to 2015, with an average annual increase of 4%. However, there were fluctuations throughout this period. Production reached a high of USD 501.000 in 2014 but fell significantly the following year. This decline is reflected in the export figures 2015, which showed a substantial decrease compared to 2014, reaching USD 577.000. This represented a 35% decline from 2007.

Figure 1. Distribution of industrial and non-industrial production of edible mushrooms in Ecuador.

In contrast, consumption of dried mushrooms rose considerably over the same period. In 2014, consumption peaked at USD 889.000 before falling dramatically in 2015 in terms of market value (USD 576.900) and production (25 tons). On the contrary, in 2007-2015, overall consumption steadily increased. Consumption reached a record high of 35 tons in 2014 before significantly dipping the following year. Despite a notable decline in revenue in 2015, the Ecuadorian dried mushroom industry experienced a growth in consumption leading up to that year ⁴⁵.

According to a market analysis conducted in Quito to evaluate the potential selling of P. ostreatus as a source of fungal protein, it has been observed that people most interested in consuming edible mushrooms are those who take care of their physical health through good dietary practices and people with socio-environmental awareness. According to the National Institute of Statistics and Censuses of Ecuador, mushrooms are not considered in the basic food basket, costing USD 712.11 (January 2021). Thus, edible mushrooms are intended to be offered to consumers with an income above the basic food basket ⁴⁶.

Moreover, in a recent study in Los Rios province of Ecuador, approx. The 77% of people involved in a study about mycophagy's state of the art indicated their interest in cultivating mushrooms ⁴⁷. A similar scenario was recorded in Cuenca (the regional capital city of the Andean province of Azuay), where 82% of inhabitants, aged between 15- and 74 years old, used to consume edible mushrooms in their diet ⁴⁸. Consumers' growing desire for healthy, eco-friendly products has driven demand for locally produced alternatives. This shift in dietary preferences presents a commercial opportunity for Ecuador to expand its cultivation of edible mushrooms ⁴⁹.

BIOTECHNOLOGICAL USE OF AGRICULTURAL BY-PRODUCTS IN ECUADOR: PRODUCTION OF EDIBLE MUSHROOMS

Edible mushroom cultivation on lignocellulosic substrates dates back hundreds of years ⁵⁰. Substrate features influence cultivated mushrooms' chemical, functional, and sensory characteristics. Substrate decomposition is related to fungi's ability to secrete hydrolytic enzymes (cellulases, xylanases and tannases) and oxidase enzymes that degrade plant by-products ⁵². Design and control of edible mushroom production is of great importance, as well as having food that preserves its nutritional value and is microbiologically safe. This process usually occurs indoors in small greenhouses and involves solid-state fermentation. This biomass can include different types of by-products from agro-industrial production ⁵². From this perspective, coculture has become a prominent alternative. Fungi used in coculture are saprophytes, and thanks to lignocellulosic exoenzyme production, they act as converters of agricultural and forest waste biomass into carpophores, which are exploited as food ⁴⁰. The process includes several operations: inoculum preparation, substrate preparation, inoculation, production, and harvest. The inoculum consists of pure mycelium of a specific strain obtained from spores, mushroom pieces, or germplasm sell providers. The collected mycelium is transferred to cereal grains such as wheat, rice or rye to produce the propagation material known as “spawn.” The substrate is previously selected according to particle size and the presence of pathology symptoms. The next step is pasteurization, which reduces biological contaminants that can disturb mushroom growth; it usually happens at 60 °C for 1-2 hours. After substrate cooling, inoculation is carried out using the spawn previously prepared by distributing the inoculum homogeneously into the cultivation substrate. Depending on the mushroom species, the cultivation techniques differ ⁵³.

Agaricus mushrooms are grown on trays, using compost as the growing medium. The process happens in tunnels, which are unique fermentation rooms. Edible mushrooms grow on the compost surface and are cut and harvested at the mature stage. Lentinus is traditionally grown on wood logs, but in Ecuador, it has been replaced by artificial log cultivation, utilizing heat-treated substrates composed of sawdust enclosed in plastic bags, known as “bag-log cultivation.” This approach offers shorter crop cycles and increased yields. Pleurotus is cultivated in polyethylene bags arranged in stacks on shelves or bag-suspended systems hooked on the ceiling. For Lentinus and Pleurotus production, after the incubation time, perforations on the bag edges are made to allow carpophores to grow outside and be harvested. Enhancing productivity in mushroom cultivation requires assessing a control system that monitors the environmental parameters in the growing rooms, such as temperature, relative humidity, airflow, oxygen and carbon dioxide content. In Ecuador, the harvest is made manually, and mushrooms are sold chiefly as fresh food; only a few producers transform them into pickled products ⁵⁴.

Mushroom cultivation is a valuable alternative to use agricultural wastes and thus to reduce environmental pollution ⁵. Many different types of agro-industrial wastes are used, both of plant and animal origin (Figure 2) ^51,55-59.

Studies showed that mixing agro-industrial wastes can increase biological efficiency and Mushroom production compared to single-substrate cultivation. Different substrates have different cellulose, hemicellulose and lignin contents that contribute to the nutritional requirements of growing fungi ^60,61. In the Ecuadorian economic context, edible mushroom cultivation may become a sustainable strategy for the use of agricultural waste. The country's economy is mainly based on agricultural production, which generates large amounts of waste ⁶². The most significant agricultural production is focused on permanent crops such as cocoa (Theobroma cacao), African palm (Elaeis guineensis), banana (Musa paradisiaca), sugar cane (Saccharum officinarum) and is spread over 1.42 million hectares. The cocoa area represents 41.8% of the total permanent crop area, implying a high production of by-products. Studies have shown that 80% of fruit is discarded, giving rise to by-products such as seed husk and fruit mesocarp ⁴⁰. Sugar cane (S. officinarum) is a relevant crop, too. The juice extraction process generates bagasse, the fibrous residual part of the stalk, as a by-product. Cellulose, hemicellulose, lignin, ash, and wax characterize its chemical composition. Bagasse represents approximately 30% of the plant's weight and is one of the most abundant agro-industrial wastes in the world, with an annual production of 1900 million tons. In Ecuador, approximately 8.6 million tonnes are produced annually, ^63,64. It is a versatile material that produces paper, biofuel, and animal feed. Due to its chemical composition and texture, it lends itself as a substrate for solid-state fermentation processes ⁶³, such as mushroom production.

Figure 2. Growing substrates available in Ecuador for edible mushroom production are obtained from plant and animal waste.

Coconut palm (Cocos nucifera) is another permanent crop that is highly present in Ecuador. In 2019, total coconut production was 6.2 metric tons, with Manabí province contributing 1.8 metric tons. Farmers in Manabí use palm residues and trunk fiber as compost, while coconuts sold by small-scale traders are incinerated. In Manta, coconut waste generated by merchants and consumers is disposed of in municipal landfills, generating waste management problems ^65,66.

The availability of agricultural by-products is an essential factor in planning the production of edible mushrooms; however, it is pivotal to consider their safety for human health. Studies have shown that P. ostreatus, A. bisporus, and H. ercinaceus mushrooms grown on mercury-containing substrates have accumulated mercury in the carpophore and stem without altering the composition of macronutrients such as carbohydrates, proteins, and fats. These results demonstrate the fundamental role of selecting suitable growing substrates that avoid exposure to mercury levels harmful to human health ⁶⁷.

Regarding economic feasibility, edible mushroom production in Ecuador using agricultural wastes recorded a lack of official data from governmental institutions, such as production costs and market prices. Experimental studies on local universities in the Azuay province showed that the production costs of fresh mushrooms fluctuated between USD 4.2/kg ⁶⁸ to 6.8/kg⁴⁸, while market prices varied between USD 10/kg ⁴⁸ to 26/kg ⁶⁹. These costs and prices mainly varied depending on the mushroom species and infrastructure availability. The Ecuadorian Ministry of Agriculture and Livestock (MAGAP) reported specific mushroom production experiences in the Amazonian region ⁷⁰. However, the Ecuadorian government considers the rural and agricultural sector a key driver of economic growth, prioritizing food production and developing long-term, sustainable activities that create jobs and income ⁷¹. Government and non-governmental incentives are available for farmers who want to invest in small agriculture production, such as the Agrobiodiversity, Seed and Sustainable Agriculture Research Fund (FIASA) ⁷² and the Inter-American Development Bank (IDB) ⁷³. With these premises, edible mushroom production could become an opportunity for economic development in Ecuador, even in rural areas.

CONCLUSIONS

Cultivated edible mushrooms are a source of protein low in fat and calories. Secondary metabolites such as polyphenols, lactones and other bioactive compounds increase their commercial value and interest in customers. Cultivated mushrooms are saprophytes; they grow on organic decomposing substrates, such as waste from agriculture and the paper industry. Ecuador is a country based on agriculture. In fact, after oil, agricultural exports like bananas, cocoa, and shrimp are the country's second-largest source of income. The production process generates tons of by-products that can be repurposed as growing substrates for edible mushrooms. Ecuador's abundant plant and animal waste and the potential for creating substrate mixtures provide the technical foundation for utilizing agricultural waste through coculture.

In addition, more studies are needed on the medicinal properties of the edible mushrooms considered in this review to gain a more comprehensive knowledge of their properties.

However, it is necessary to ensure that substrates are free of chemical or physical contaminants that mushrooms could carry during production. This literature review suggests further investigation into the following aspects: a) carry out experiments on new edible mushroom species using local agro-industrial wastes, b) identify promising native species for use in the agro-industrial sector, c) carry out studies on the presence of heavy metals or other undesirable compounds.

Author Contributions: Conceptualization, Laura Scalvenzi and Amaury Pérez Martínez; methodology, Matteo Radice, Renier Abreu Naranjo and Mayra Chiriboga Ruilova; resources, Matteo Radice and Estela Guardado Yordi; data curation, Matteo Radice; investigation Amaury Pérez Martínez, Mayra Chiriboga Ruilova and Laura Scalvenzi; writing—original draft preparation Amaury Pérez Martínez, Mayra Chiriboga Ruilova and Laura Scalvenzi; writing—review and editing, Estela Guardado Yordi and Renier Abreu Naranjo; supervision, Laura Scalvenzi. All authors have read and agreed to the published version of the manuscript.

Funding: This research received no external funding. The APC was funded by the Universidad Estatal Amazónica.

Institutional Review Board Statement: Not applicable.

Informed Consent Statement: Not applicable.

Data Availability Statement: Not applicable.

Acknowledgments: The manuscript's authors thank Universidad Estatal Amazónica for the facilities provided in the research development.

Conflicts of Interest: The authors declare no conflict of interest.

REFERENCES

1. Gautam, A.K.; Verma, R.K.; Avasthi, S.; Sushma; Bohra, Y.; Devadatha, B.; Niranjan, M.; Suwannarach, N. Current Insight into Traditional and Modern Methods in Fungal Diversity Estimates. Journal of Fungi 2022, 8 (3), 226

2. Prasad, S.; Rathore, H.; Sharma, S.; Yadav, A. Medicinal mushrooms as a source of novel functional food. International Journal of Food Science, Nutrition and Dietetics 2015, 4 (5), 221-5 https://doi.org/10.19070/2326-3350-1500040:https://doi.org/10.19070/2326-3350-1500040.

3. Ortiz Rodríguez, E.J.; Pineda Insuasti, J.; Duarte Trujillo, A.S.; Parrado Castro, A.G.; Soto Arroyave, C.P.; Pineda Soto, C.A. La producción de hongos comestibles frente a la crisis alimentaria del Ecuador. Revista Biorrefinería 2018, 1 (1), 13-22

4. Martínez-Carrera, D. Mushroom biotechnology in tropical America. International Journal of Mushroom Sciences 2000, 3 (1), 9-20

5. Mihai, R. A.; Melo Heras, E.J.; Florescu, L.I.; Catana, R.D. The Edible Gray Oyster Fungi Pleurotus ostreatus (Jacq. ex Fr.) P. Kumm a Potent Waste Consumer, a Biofriendly Species with Antioxidant Activity Depending on the Growth Substrate. Journal of Fungi 2022, 8 (3), 274 https://doi.org/10.3390/jof8030274 https://doi.org/10.3390/jof8030274

6. Zhang, Y.; Geng, W.; Shen, Y.; Wang, Y.; Dai, Y.C. Edible Mushroom Cultivation for Food Security and Rural Development in China: Bio-Innovation, Technological Dissemination and Marketing. Sustainability 2014, 6 (5), 2961-2973 https://doi.org/10.3390/su6052961 https://doi.org/10.3390/su6052961

7. Silva-Neto, C.d.M. e.; Pinto, D.d.S.; Santos, L.A.C.; Cala ÇA, FJS; Almeida, S.d.S. Food production potential of Favolus brasiliensis (Basidiomycota: Polyporaceae), an indigenous food. Food Science and Technology 2021, 41 (suppl 1), 183–188 https://doi.org/10.1590/fst.12620:https://doi.org/10.1590/fst.12620.

8. Kaliyaperumal, M.; Kezo, K.; Gunaseelan, S. A Global Overview of Edible Mushrooms. In Biology of Macrofungi, 1^st ed.; Singh, B.P.; Lallawmsanga; Passari, A.K., Eds.; Springer International Publishing: Cham, 2018; pp 15-56.

9. Kumar Sharma, S.; Gautam, N., Chemical and Bioactive Profiling, and Biological Activities of Coral Fungi from Northwestern Himalayas. Scientific Reports 2017, 7 (1), 46570 https://doi.org/10.1038/srep46570:https://doi.org/10.1038/srep46570.

10. Khumlianlal, J.; Sharma, K.C.; Singh, L.M.; Mukherjee, P.K.; Indira, S. Nutritional Profiling and Antioxidant Property of Three Wild Edible Mushrooms from North East India. Molecules 2022, 27 (17), 5423 https://doi.org/10.3390/molecules27175423 https://doi.org/10.3390/molecules27175423

11. Thu, Z. M.; Myo, KK; Aung, H.T.; Clericuzio, M.; Armijos, C.; Vidari, G. Bioactive Phytochemical Constituents of Wild Edible Mushrooms from Southeast Asia. Molecules 2020, 25 (8), 1972 https://doi.org/10.3390/molecules25081972 https://doi.org/10.3390/molecules25081972

12. Muszyńska, B.; Grzywacz-Kisielewska, A.; Kała, K.; Gdula-Argasińska, J. Anti-inflammatory properties of edible mushrooms: A review. Food Chemistry 2018, 243, 373-381 https://doi.org/10.1016/j.foodchem.2017.09.149:https://doi.org/10.1016/j.foodchem.2017.09.149.

13. Fijałkowska, A.; Jędrejko, K.; Sułkowska-Ziaja, K.; Ziaja, M.; Kała, K.; Muszyńska, B. Edible Mushrooms as a Potential Component of Dietary Interventions for Major Depressive Disorder. Foods 2022, 11 (10), 1489 https://doi.org/10.3390/foods11101489 https://doi.org/10.3390/foods11101489

14. Mustafa, F.; Chopra, H.; Baig, A.A.; Avula, S.K.; Kumari, S.; Mohanta, T.K.; Saravanan, M.; Mishra, A.K.; Sharma, N.; Mohanta, Y.K. Edible Mushrooms as Novel Myco-Therapeutics: Effects on Lipid Level, Obesity and BMI. Journal of Fungi 2022, 8 (2), 211 https://doi.org/10.3390/jof8020211 https://doi.org/10.3390/jof8020211

15. Das, A.; Chen, C.M.; Mu, SC; Yang, S.H.; Ju, Y.M.; Li, SC Medicinal Components in Edible Mushrooms on Diabetes Mellitus Treatment. Pharmaceutics 2022, 14 (2), 436 https://doi.org/10.3390/pharmaceutics14020436 https://doi.org/10.3390/pharmaceutics14020436

16. Seo, D. J.; Choi, C.,Antiviral Bioactive Compounds of Mushrooms and Their Antiviral Mechanisms: A Review. Viruses 2021, 13 (2), 350 https://doi.org/10.3390/v13020350:https://doi.org/10.3390/v13020350.

17. Assemie, A.; Abaya, G. The Effect of Edible Mushroom on Health and Their Biochemistry. International Journal of Microbiology 2022, 2022, 8744788 https://doi.org/10.1155/2022/8744788:https://doi.org/10.1155/2022/8744788.

18. Ibrahim, R.M.; Ali, M.I.K.; Abdel-salam, F.F. Nutritional and Quality Characteristics of Some Foods Fortified with Dried Mushroom Powder as a Source of Vitamin D. International Journal of Food Science 2022, 2022, 2792084 https://doi.org/10.1155/2022/2792084:https://doi.org/10.1155/2022/2792084.

19 Sganzerla, W.G.; Todorov, S.D.; da Silva, A.P.G., Research Trends in the Study of Edible Mushrooms: Nutritional Properties and Health Benefits. International Journal of Medicinal Mushrooms 2022, 24 (5), 1 – 18

20 Anusiya, G.; Gowthama Prabua, U.; Yamini, N.V.; Sivarajasekar, N.; Rambabu, K.; Bharath, G.; Banat, F. A review of the therapeutic and biological effects of edible and wild mushrooms. Bioengineered 2021, 12 (2), 11239–11268 https://doi.org/10.1080/21655979.2021.2001183

21 Kumar, K.; Mehra, R.; Guiné, R.P.F.; Lima, M.J.; Kumar, N.; Kaushik, R.; Ahmed, N.; Yadav, A.N.; Kumar, H. Edible Mushrooms: A Comprehensive Review on Bioactive Compounds with Health Beneﬁts and Processing Aspects. Foods 2021, 10, 2996. https://doi.org/10.3390/foods10122996

22 Ma, G.; Du, H.; Hu, Q.; Yang, W.; Pei, F.; Xiao, H. Health benefits of edible mushroom polysaccharides and associated gut microbiota regulation. Critical Reviews in Food Science and Nutrition 2021, 1-18 DOI: 10.1080/10408398.2021.1903385

23 Jayachandran, M.; Xiao, J.; Xu, B. A Critical Review on Health Promoting Beneﬁts of Edible Mushrooms through Gut Microbiota. International Journal of Molecular Sciences 2017, 18, 1934.

24 Zhao, Q.; Jiang ,Y.; Zhao, Q.; Patrick Manzi, H.; Su, L.; Liu, D.; Huang, X.; Long, D.; Tang, Z.; Zhang, Y. The beneﬁts of edible mushroom polysaccharides for health and their inﬂuence on gut microbiota: a review. Frontiers in Nutrition 2023, 10: 1213010. doi: 10.3389/fnut.2023.121301025

25 Cerletti, C.; Esposito, S.; Iacoviello, L. Edible Mushrooms and Beta-Glucans: Impact on Human Health. Nutrients 2021, 13, 2195. https://doi.org/10.3390/nu13072195

26 Phan, C. W.; David, P.; Sabaratnam, V. Edible and Medicinal Mushrooms: Emerging Brain Food for the Mitigation of Neurodegenerative Diseases. Journal od Medicinal Food 2017, 20 (1), 1–10 doi: 10.1089/jmf.2016.3740

27 Sabaratnam, V.; Kah-Hui, W.; Naidu, M.; David, P.R. Neuronal Health – Can Culinary and Medicinal Mushrooms Help? Journal of Traditional and Complementary Medicine 2011, 3 (1), 62-68

28. Guillamón, E.; García-Lafuente, A.; Lozano, M.; D´Arrigo, M.; Rostagno, M.A.; Villares, A.; Martínez, J.A. Edible mushrooms: Role in the prevention of cardiovascular diseases. Fitoterapia 2010, 81 (7), 715-723 https://doi.org/10.1016/j.fitote.2010.06.005:https://doi.org/10.1016/j.fitote.2010.06.005.

29. Mattila, P.; Könkö, K.; Eurola, M.; Pihlava, J.M.; Astola, J.; Vahteristo, L.; Hietaniemi, V.; Kumpulainen, J.; Valtonen, M.; Piironen, V. Contents of Vitamins, Mineral Elements, and Some Phenolic Compounds in Cultivated Mushrooms. Journal of Agricultural and Food Chemistry 2001, 49 (5), 2343-2348 https://doi.org/10.1021/jf001525d:https://doi.org/10.1021/jf001525d.

30. Reis, F. S.; Barros, L.; Martins, A.; Ferreira, ICFR Chemical composition and nutritional value of the most widely appreciated cultivated mushrooms: An inter-species comparative study. Food and Chemical Toxicology 2012, 50 (2), 191-197 https://doi.org/10.1016/j.fct.2011.10.056:https://doi.org/10.1016/j.fct.2011.10.056.

31. Atila, F., Nadhim; Owaid, M. N.; Shariati, M.A. The nutritional and medical benefits of Agaricus bisporus: a review. Journal of Microbiology, Biotechnology and Food Sciences 2017, 7 (3), 281–286 https://doi.org/10.15414/jmbfs.2017/18.7.3.281-286:https://doi.org/10.15414/jmbfs.2017/18.7.3.281-286.

32. Carneiro, A.A.J.; Ferreira, ICFR; Dueñas, M.; Barros, L.; da Silva, R.; Gomes, E.; Santos-Buelga, C. Chemical composition and antioxidant activity of dried powder formulations of Agaricus blazei and Lentinus edodes. Food Chemistry 2013, 138 (4), 2168-2173 https://doi.org/10.1016/j.foodchem.2012.12.036:https://doi.org/10.1016/j.foodchem.2012.12.036.

33. Hassan, E.M.; El Gendy, A.E.N.G.; Abd-ElGawad, A.M.; Elshamy, A.I.; Farag, M.A.; Alamery, S.F.; Omer, E.A. Comparative Chemical Profiles of the Essential Oils from Different Varieties of Psidium guajava L. Molecules 2021, 26 (1), 119 https://doi.org/10.3390/molecules26010119 https://doi.org/10.3390/molecules26010119

34. Pardo-Giménez, A.; Figueirêdo, V.; Dias, E. S.; Pardo-González, J. E.; Álvarez-Ortí, M.; Zied, D. C. Análisis proximal de carpóforos de Agaricus subrufescens Peck producidos sobre diferentes capas de cobertura. ITEA Producción Vegetal 2013, 109 (3), 290-302 https://doi.org/10.12706/itea.2013.017:https://doi.org/10.12706/itea.2013.017.

35. Vinhal Costa Orsine, J.; Carvalho Garbi Novaes, M.; Ramírez Asquieri, E. El valor nutritivo de Agaricus sylvaticus: setas cultivadas en Brasil. Nutrición Hospitalaria 2012, 27 (2), 449-455 https://doi.org/10.3305/nh.2012.27.2.5504:https://doi.org/10.3305/nh.2012.27.2.5504.

36. Smiderle, F.R.; Carbonero, E.R.; Sassaki, G.L.; Gorin, P.A.J.; Iacomini, M. Characterization of a heterogalactan: Some nutritional values of the edible Mushroom Flammulina velutipes. Food Chemistry 2008, 108 (1), 329-333 https://doi.org/10.1016/j.foodchem.2007.10.029:https://doi.org/10.1016/j.foodchem.2007.10.029.

37. Bach, F.; Helm, C.; De Lima, E.A.; Barba Bellettini, M.; Haminiuk, C.W.I. Influence of cultivation methods on the chemical and nutritional characteristics of Lentinula edodes. Emirates Journal of Food and Agriculture 2019, 30 (12), 1006-1013 https://doi.org/10.9755/ejfa.2018.v30.i12.1879 https://doi.org/10.9755/ejfa.2018.v30.i12.1879

38. Raman, J.; Jang, K.Y.; Oh, Y.L.; Oh, M.; Im, J.H.; Lakshmanan, H.; Sabaratnam, V. Cultivation and Nutritional Value of Prominent Pleurotus spp.: An Overview. Mycobiology 2021, 49 (1), 1-14 https://doi.org/10.1080/12298093.2020.1835142:https://doi.org/10.1080/12298093.2020.1835142.

39. Vega, A.; Franco, H. Productividad y calidad de los cuerpos fructíferos de los hongos comestibles Pleurotus pulmonarius RN2 y P. djamor RN81 y RN82 cultivados sobre sustratos lignocelulósicos. Información tecnológica 2013, 24 (1), 69-78 http://dx.doi.org/10.4067/S0718-07642013000100009.:http://dx.doi.org/10.4067/S0718-07642013000100009.

40.Vallejo Torres, C.; Díaz Ocampo, R.; Morales Rodríguez, W.; Vera Chang, J.; Cortéz Salazar, T. Calidad alimenticia del hongo Pleurotus ostreatus, fresco y deshidratado, cultivado en tres residuos agrícolas. Revista ESPAMCIENCIA 2017, 8 (2), 75-83

41. Gamboa-Trujillo, J. P.; Wartchow, F.; Cerón, C.; Aules, E.; Aigaje, C.; Calvalcant, i. LH; Gibertoni, T.B. Traditional use of Gymnopus nubicola as food resource in a Kichwa community, Pichincha, Ecuador. Mycosphere 2014, 5 (1), 180–186 https://doi.org/10.5943/mycosphere/5/1/9:https://doi.org/10.5943/mycosphere/5/1/9.

42. Gamboa-Trujillo, P.; Wartchow, F.; Cerón-Martinez, C.; Andi, D.; Uwinjin, P.; Grefa, G.; Entza, M.; Chimbo, E.; Chimbo, J.; Payaguaje, J.; Piyaguaje, N.; Payaguaje, D.; Licuy, V.; López, V.; Mendua, M.; Criollo, M.; Jati, M.; De La Cruz, S.; Calazacón, M.; Flores, S.; Aules, E.; Aigaje, C.; De Aro, M.; Morales, A.; Murillo, Y.; Farinango, G.; Gibertoni, T. Edible Mushrooms of Ecuador: consumption, myths and implications for conservation. Ethnobotany Research and Applications 2019, 18 (0), 1-15 http://dx.doi.org/10.32859/era.18.38.1-15:http://dx.doi.org/10.32859/era.18.38.1-15.

43. Industrialización de hongos comestibles. Secretaria General de la Organización de los Estados Americanos. Available online: https://www.oas.org/DSD/publications/Unit/oea60s/ch20.htm#TopOfPage (accessed on 10 May 2024).

44. Calero Guevara, L.E. Valoración del crecimiento del hongo ostra rosado (Pleurotus djamor) sobre formulaciones de sustra-tos de residuos agroindustriales y forestales de la provincia de Cotopaxi para la producción de setas comestibles en la empres ASOPROTEC. Universidad Técnica de Ambato, Ecuador, 2018.

45 Dried Mushroom's market in Ecuador, IndexBox. Available online: https://app.indexbox.io/report/071231h071232h071233h071239/218/ (accessed 10 May 2024);

46 Instituo Nacional de Estadística y Censos. Quito (Ecuador). Boletín técnico N. 01-2021-IPC-Indices del precios al consumidor. Available online: https://www.ecuadorencifras.gob.ec/documentos/web-inec/Inflacion/2021/Enero-2021/Boletin_tecnico_01-2021-IPC.pdf (accessed 10 May 2024)

47 Reyes Borja, W.O.; Romero, M.E.; Arana Vera, L.P.; H. Duran, P.H.; Reyes. RG State of the art of mycophagy in Los Rios, Ecuador. Advances in Environmental Biology 2016, 10 (4)

48 Uguña Naula, P.X. Producción y comercialización del hongo comestible denominado champiñon en el cantón Cuenca, provincia Azuay. Master thesis, Universidad Politécnica Salesiana, Quito (Ecuador), 2021

49. Padilla Lituma, S. C. El análisis del mercado potencial para el hongo Pleurotus ostreatus como proteína orgánica y fuente nutricional para el consumo humano en el distrito metropolitano de Quito. TDegree thesis, Universidad de las Fuerzas Armadas, Quito (Ecuador), 2020

50. Chen, F.; Grimm, A.; Eilertsen, L.; Martín, C.; Arshadi, M.; Xiong, S. Integrated production of edible Mushroom (Auricularia auricular-judae), fermentable sugar and solid biofuel. Renewable Energy 2021, 170, 172-180 https://doi.org/10.1016/j.renene.2021.01.124:https://doi.org/10.1016/j.renene.2021.01.124.

51. Bellettini, M.B.; Fiorda, F.A.; Maieves, H.A.; Teixeira, G. L.; Ávila, S.; Hornung, P. S.; Júnior, A.M.; Ribani, R.H. Factors affecting mushroom Pleurotus spp. Saudi Journal of Biological Sciences 2019, 26 (4), 633-646 https://doi.org/10.1016/j.sjbs.2016.12.005:https://doi.org/10.1016/j.sjbs.2016.12.005.

52. Díaz Muñoz, K.; Casanova Guajardo, M.; León Torres, C.A.; Gil Ramírez, L.A.; Bardales Vásquez, C.B.; Cabos Sánchez, J. Producción de Pleurotus ostreatus (Pleurotaceae) ICFC 153/99 cultivado sobre diferentes residuos lignocelulósicos. Arnaldoa 2019, 26 (3), 1177-1184 http://dx.doi.org/10.22497/arnaldoa.263.26322:http://dx.doi.org/10.22497/arnaldoa.263.26322.

53. Melo de Carvalho, C.S.; Sales-Campos, C.; Nogueira de Andrade, M.C. Mushroom of the Pleurotus genus: a review of cultivation techniques. Interciencia 2010, 35 (3), 177-182

54. Sanchez, C. Cultivation of Pleurotus ostreatus and other edible mushrooms. Appl. Microbiol Biotechnol 2010, 85: 1321-1337. Doi: 10.1007/s00253-009-2343-7

55. Zervakis, G.; Philippoussis, A.; Ioannidou, S.; Diamantopoulou, P. Mycelium growth kinetics and optimal temperature conditions for the cultivation of edible mushroom species on lignocellulosic substrates. Folia Microbiologica 2001, 46 (3), 231-234 https://doi.org/10.1007/BF02818539:https://doi.org/10.1007/BF02818539.

56. Andrino, A.; Morte Gómez, M.A.; Honrubia García, M. In Caracterización y cultivo de tres cepas de Pleurotus eryngii (Fries) Quélet sobre sustratos basados en residuos agroalimentarios, Anales de Biología, Murcia: Universidad de Murcia, Servicio de Publicaciones: 2011.

57. Pineda-Insuasti, J.A.; Ramos-Sánchez, L.B.; Soto-Arroyave, C.P. Producción de Pleurotus ostreatus por fermentación en estado sólido: una revisión. ICIDCA. Sobre los Derivados de la Caña de Azúcar 2014, 48 (2), 13-23

58. Win, T.T.; Ohga, S. Study on the cultivation of Agaricus blazei (Almond Mushroom) grown on compost mixed with selected agro-residues. Advances in Microbiology 2018, 8 (10), 778-789 https://doi.org/10.4236/aim.2018.810051:https://doi.org/10.4236/aim.2018.810051.

59. Otieno, OD; Mulaa, F.J.; Obiero, G.; Midiwo, J. Utilization of fruit waste substrates in mushroom production and manipulation of chemical composition. Biocatalysis and Agricultural Biotechnology 2022, 39, 102250 https://doi.org/10.1016/j.bcab.2021.102250:https://doi.org/10.1016/j.bcab.2021.102250.

60. Kuhad, R.C.; Singh, A.; Eriksson, K.E.L. Microorganisms and enzymes involved in the degradation of plant fiber cell walls. In Biotechnology in the Pulp and Paper Industry, Eriksson, K.E.L.; Babel, W.; Blanch, H.W.; Cooney, C.L.; Enfors, S.O.; Eriksson, K.E.L.; Fiechter, A.; Klibanov, A.M.; Mattiasson, B.; Primrose, S.B.; Rehm, H.J.; Rogers, P.L.; Sahm, H.; Schügerl, K.; Tsao, G.T.; Venkat, K.; Villadsen, J.; von Stockar, U.; Wandrey, C., Eds. Springer Berlin Heidelberg: Berlin, Heidelberg, 1997; pp 45-125.

61. Owaid, M.N.; Abed, A.M.; Nassar, B.M. Recycling cardboard wastes to produce blue oyster mushroom Pleurotus ostreatus in Iraq. Emirates Journal of Food and Agriculture 2015, 27 (07), 537-541 https://doi.org/10.9755/ejfa.2015.04.118:https://doi.org/10.9755/ejfa.2015.04.118.

62. Buendía-Martínez, I.; Álvarez-Herranz, A.; Moreira Menéndez, M. Business Cycle, SSE Policy, and Cooperatives: The Case of Ecuador. Sustainability 2020, 12 (13), 5485

63. Loh, Y.R.; Sujan, D.; Rahman, M.E.; Das, C.A. Sugarcane bagasse—The future composite material: A literature review. Resources, Conservation and Recycling 2013, 75, 14-22 https://doi.org/10.1016/j.resconrec.2013.03.002:https://doi.org/10.1016/j.resconrec.2013.03.002.

64. Peñafiel, M.E.; Matesanz, J.M.; Vanegas, E.; Bermejo, D.; Mosteo, R.; Ormad, M.P. Comparative adsorption of ciprofloxacin on sugarcane bagasse from Ecuador and on commercial powdered activated carbon. Science of The Total Environment 2021, 750, 141498 https://doi.org/10.1016/j.scitotenv.2020.141498:https://doi.org/10.1016/j.scitotenv.2020.141498.

65. San Adrés Zevallos, G.I.; Aguilar Sierra, S.M., Estado actual del ciclo de vida del coco en Manabí, Ecuador. VIII Congreso Internacional de Investigación REDU, 25 abril 2022, http://doi.org/10.5867/Medwave.2022.S1.CI37

66. Mihai, R.A.; Melo Heras, E.J.; Florescu, F.; Catana, R.D. The Edible Gray Oyster Fungi Pleurotus ostreatus (Jacq. ex Fr.) P. Kumm a Potent Waste Consumer, a Biofriendly Species with Antioxidant Activity Depending on the Growth Substrate. Journal of Fungi 2022, 8 (3), 274; https://doi.org/10.3390/jof8030274

67. Rzymski, P.; Mleczek, M.; Siwulski, M.; Gąsecka, M.; Niedzielski, P. The risk of high mercury accumulation in edible mushrooms cultivated on contaminated substrates. Journal of Food Composition and Analysis 2016, 51, 55-60 https://doi.org/10.1016/j.jfca.2016.06.009:https://doi.org/10.1016/j.jfca.2016.06.009.

68. Mogrovejo Lazo, A.; Arias Abad, D.; Vidal Astudillo, B., Producción y Comercialización de champiñones en la provincia de Azuay-Ecuador. Estudio de Factibilidad. Telos: revista de Estudios Interdisciplinarios en Ciencias Sociales 2020, 22 (1), 144-161; DOI: www.doi.org/10.36390/telos221.10

69. Selina Wamucii. Ecuador mushrooms and truffle prices [Internet]. Nairobi 2024, [cited 2024 May 13]. Available from: https://www.selinawamucii.com/insights/prices/ecuador/mushrooms-and-truffles/#wholesale-prices.

70. The Ministry of Agriculture and Livestock of Ecuador. Productores de Pastaza participan de proyecto piloto en cultivo de hongos ostras. Available online: https://www.agricultura.gob.ec/productores-de-pastaza-participan-de-proyecto-piloto-en-cultivo-de-hongos-ostras/ (Accessed on 13 May 2024)

71. International Fund for Agriculture Development-IFAD. Country Strategy Note – Ecuador 22-24. Latin America and Carribbean Division, Programme Management Department. Quito. 2022; Available online: https://www.ifad.org/documents/38711624/39485439/CSN-Ecuador-22-24.pdf/f94890c1-b962-7106-5544-c1d77a1ff90b?t=1710238587690 (accessed 13 May 2024);

72. Gobierno del Ecuador. Gobierno del Encuentro-Guillermo Lasso. Preasignados proforma, Pag. 2-5. Available from: https://test-assets opsaa.iica.int/storage/initiative/2024/03/25998155494b1508483e2ac947a7c8e0.pdf (accessed 13 May 2024).

73. Inter-American Development Bank (IDB). Ecuador to expand financing and investment for Amazonian Bio-businesses, with IDB Support. Available online: https://www.iadb.org/en/news/ecuador-expand-financing-and-investment-amazonian-bio-businesses-idb-support (accessed 13 May 2024)

Received: December 14, 2023/ Accepted: February 25, 2024 / Published: June 15, 2024

Citation: Scalvenzi L, Chiriboga Ruilova M V, Guardado Yordi E, Radice M,

Abreu Naranjo R, Pérez Maertínez A. Edible mushroom production in Ecuador: opportunities for biotechnological use of agricultural by-products.Bionatura journal 2024; 1 (2) 15. http://dx.doi.org/10.21931/BJ/2024.01.02.15

Additional information Correspondence should be addressed to amperez@uea.edu.ec

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