A Systematic Review of the Seven Most Cultivated Mushrooms: Production Processes, Nutritional Value, Bioactive Properties and Impact on Non-Communicable Diseases

Abstract

Mushroom production, mushroom knowledge and mushroom cultivation have aroused the interest of many researchers, scientists, institutions, cultural and mushroom-loving associations, and ordinary mushroom pickers as well. The contribution of wild mushrooms to the economic, cultural and touristic development and social cohesion of the Greek rural population is considered very important. In addition to their therapeutic and dietary value, they are also valuable for their diverse chemical and medicinal properties. The PubMed—Medline, Web of Science and Cochrane Library databases were searched for relevant articles published up to January, 2025. Even today, some macromycetes are still used in medicine due to their remarkable healing properties. In the form of powders or solutions, they were formerly used as a remedy for epilepsy, tuberculosis, nervous diseases and various severe inflammations. This review documents the benefits of mushroom consumption suggested by health experts for pathological conditions and health improvement and highlights their superiority as non-animal protein sources according to their nutrients.

1. Introduction

Non-communicable diseases (NCDs) such as cardiovascular diseases, diabetes, neurological disorders and cancer are responsible for 80% of diseases in the European Union [1]. Nutrition is the key to many of these diseases, not only to reduce the increasing incidence but also to limit the consequences, such as hyperglycemia, hypocalcemia, hypertension, etc. [2]. Consumers tend to look for functional foods as alternative medical solutions [1]. Cultivated mushrooms could be an environmentally friendly solution with benefits for the planet due to their cultivation method, but also a sustainable nutritious food [3].

The global market for mushrooms is worth USD 63 billion, with cultivated edible mushrooms accounting for 54% of this market, medicinal mushrooms for 38% and wild mushrooms for 8% [4]. The most commonly cultivated mushrooms worldwide are Agaricus bisporus, Lentinula edodes, Pleurotus spp., Auricularia auricula-judea, Volvariella volvacea and Flammulina velutipes [5], and truffles, especially black and white, are the new trend [6]. The best-known wild mushrooms, however, are Boletus edulis, Cantharellus spp., Craterellus cornucopioides, Morchella spp. and Marasmius oreades [7].

The mushroom has managed to be one of the most delicious foods from the time of gatherers and hunters to modern man. In addition to indigenous mushrooms, there are also mushrooms cultivated in intensive or extensive production units that offer everyone the opportunity to acquire what is rightly called “food medicine” [3].

They enrich the body with vitamins and amino acids of high biological value, which protect against cholesterol, anemia, sugar, arteriosclerosis, heart muscle abnormalities and cancer [8]. They are also considered a dietary food, as one kilogram of mushrooms contains only 190 calories and can replace other fattening foods while providing a feeling of satiety. All the ingredients of the mushroom are retained after cooking, except for the minerals and vitamins such as thiamine, niacin and folate [1,3].

There is no doubt that wild mushrooms have a unique nutritional profile, but due to their limited use, cultivated mushrooms are entering the global market and could serve as an important alternative medicinal source of bioactive compounds due to their nutritional profile [4]. Some reasons for the limitations on the use of wild mushrooms are toxicity risks, identification difficulties and seasonality. Additionally, new post-harvest techniques such as drying, freezing and cooking, or even storage (12 °C/12 days), guarantee the extraction of higher bioactive compounds from cultivated mushrooms and thus a higher medicinal value [1,3]. Mushrooms have over 100 therapeutic activities, with primary medicinal uses being high in antiparasitic, antibacterial, antioxidant, anti-inflammatory, anticancer, antitumor, anticoagulant, cytotoxic, hypolipidemic, antithrombotic, hypocholesterolemic, anti-HIV, antidiabetic and hepatoprotective compounds [3]. The current state of research in this area of agricultural technology is limited. However, despite the studies that have been carried out for years, there are difficulties that, together with the ignorance and prejudices of the population, represent a serious obstacle to the study, cultivation and use of mushrooms in the entire Western Hemisphere.

To date, there are only a few studies dealing with post-harvest techniques used to improve the nutritional profile of mushrooms in combination with their specific effects on non-commercial diseases. This review aims to assess the production of the seven most commonly grown mushrooms as an environmentally friendly solution for an alternative protein and nutrient source, the post-harvest methods used to improve their nutrient profile, and, as a consequence, their effect on overall health.

2. Method

2.1. Systematic Review Aim and Strategy

This systematic review was performed according to the Preferred Reporting Items for Systematic Reviews (PRISMA) 2020 guidelines and checklist [9]. This systematic review focuses on an evaluation of the production techniques and post-harvest methods in order to improve their nutrient profile without loss of their organoleptic characteristics and examines their protein content and their impact on overall health (Figure 1). A complete checklist, according to the PRISMA statement, is reported in Appendix A.

2.2. Literature Search, Study Selection, Eligibility Criteria and Quality Assessment

The search strategy included articles published from January 2010 to January 2025 in a comprehensive systematic review of the databases PubMed and Web of Science; searches used MeSH terms, specifically targeting “#Mushrooms”, “#Cultivation”, “#Post-harvest techniques”, “#Non-communicable diseases”, “#Nutritional profile”, “#Nutrients”, and “#Bioactive compounds”. The aim was to uncover the relationship between the nutritional profile of different mushroom species, with a focus on macronutrients and protein content in particular.

The search in PubMed, Web of Science, Scopus and the Cochrane Library focused on identifying articles on the topics “#Mushrooms”, “#Cultivation”, “#Post-harvest techniques“, ”#Non-communicable diseases”, “#Health benefits”, “#Cardiovascular Disease”, “#Diabetes Mellitus”, “#Cancer”, “#Hypertension”, “#Dyslipidemia”, “#Neurological Disorders”, “#Nutrition” and “#Diet”. Additional keywords such as “#Food” and “#Nutrition” were included.

Scopus and Cochrane Library searches focused on “#Mushrooms”,” “#Cultivation”, “#Post-harvest techniques”, “#Diet”, “#Nutrition”, “#Side Effects” and “#Evidence-Based Interventions”, with an additional keyword “#Food”. Abstracts were screened using Rayyan. If there was any doubt about the inclusion of an article, this was included for full-text review. Full articles were screened independently by two authors using Rayyan. Disagreements were resolved by returning to the original article along with a third senior author (O.G.), when required.

One investigator (M.D.) screened all of the included studies for risk of bias, with a separate investigator (I.C.) independently validating the risk of bias. The Cochrane (version 7.2.0) risk of bias tool was used for randomized controlled trials, and the ROBINS-I V2 (risk of bias in non-randomized studies of interventions) tool was used for cohort studies [10,11].

These databases were selected based on their reputability, comprehensiveness and relevance to healthcare and research (

Table 1. Details of the search process and the unique contributions of each database to the study.

Database	Keywords	MeSH Terms (PubMed)	Initial Articles	Duplicates Removed	Final Articles for Analysis	Contribution to Study	Reason for Inclusion
PubMed	#Mushrooms, #Cultivation, #Post-harvest techniques, #Protein, #Vitamin D, #Amino acids, #Diet, #Nutritional profile, #Nutrients, #Bioactive compounds, #Non-communicable diseases, #Health benefits, #Cardiovascular disease, #Cancer, #Diabetes mellitus, #Dyslipidemia, #Hypertension, #Neurological disorders, #Nutrition, and #Diet.	#Mushrooms, #Cultivation, #Post-harvest techniques, #Non-communicable diseases, #Nutritional profile, #Nutrients, #Bioactive compounds and #Interventions, #Diet, #Health.	6	6	6	Provided a broad understanding of the interplay between mushroom cultivation and post-harvest techniques to improve their nutritional profile but also consumption, interventions and health benefits; MeSH terms ensured precision in the search.	Widely recognized as a premier biomedical database, often used for reviews in healthcare research.
Web of Science	#Mushrooms, #Cultivation, #Post-harvest techniques, #Diet, #Nutritional profile, #Nutrients, #Bioactive compounds, #Health benefits, #Cardiovascular disease, #Diabetes mellitus, #Hypertension, #Cancer, #Dyslipidemia, #Neurological disorders, #Nutrition, and #Diet.	N/A (Web of Science doesn’t use MeSH terms)	298	18	280	Enhanced the overall coverage of literature related to mushroom nutrients, interventions and their impact on overall health.	Covering a wide range of scientific disciplines.
Scopus	#Mushrooms, #Cultivation, #Post-harvest techniques, #Nutrition, #Health, #Side Effects and #Evidence-Based Interventions.	#Mushrooms, #Health	5	5	5	Strengthened the evidence base by focusing on bean bioactive compounds related to evidence-based interventions; MeSH terms ensured specificity.	Renowned for reviews and emphasis on evidence-based interventions in healthcare research.
Cochrane Library	#Mushrooms, #Cultivation, #Post-harvest techniques, #Nutrition, #Health, #Side Effects and #Evidence-Based Interventions.	#Mushrooms, #Nutrition, #Health, #Side effects, #Evidence-Based Interventions	6	6	6	Strengthened the evidence base by focusing on bean bioactive compounds related to evidence-based interventions; MeSH terms ensured specificity.	Renowned for reviews and emphasis on evidence-based interventions in healthcare research.

), and the inclusion and exclusion criteria are shown in

Table 2. Inclusion and exclusion criteria.

Inclusion Criteria	Exclusion Criteria
Reported in English	Does not report primary and/or secondary outcomes
Treated with techniques that have been described in detail	No Comparator Group (i.e., control or alternative dietary intervention)
In vitro and in vivo studies	Non-English language publication
Randomized Controlled trial or cohort study or reviews	Sample parameters (small sample)

. The last search occurred on 5 February 2025.

Figure 1. Prisma flow chart of the study (version 2020) is as follows: for their production process [5,6,7,12,13,14,15,16,17,18,19,20,21,22,23,24,25,26,27,28,29,30,31,32,33,34,35,36,37,38,39,40,41,42,43,44,45,46,47,48,49,50,51], focused on their nutritional profile as alternative source of protein [8,22,23,24,52,53,54,55,56,57,58,59,60,61,62,63,64,65,66,67,68,69,70,71,72,73,74,75,76,77,78], the post-harvest techniques to improve their nutritional profile [22,24,62,79,80,81,82,83,84,85] and their impact on non-communicable diseases [86,87,88,89,90,91,92,93,94,95,96,97,98,99,100,101,102,103,104,105,106,107,108,109,110,111,112,113,114,115,116]. * PubMed 6, Web of Science 298, Scopus 5, Cochrane Library 6. ** Excluded by the researchers due to sample parameters (small sample) or examined/analyzed varieties that are not included in this systematic review.

Figure 1. Prisma flow chart of the study (version 2020) is as follows: for their production process [5,6,7,12,13,14,15,16,17,18,19,20,21,22,23,24,25,26,27,28,29,30,31,32,33,34,35,36,37,38,39,40,41,42,43,44,45,46,47,48,49,50,51], focused on their nutritional profile as alternative source of protein [8,22,23,24,52,53,54,55,56,57,58,59,60,61,62,63,64,65,66,67,68,69,70,71,72,73,74,75,76,77,78], the post-harvest techniques to improve their nutritional profile [22,24,62,79,80,81,82,83,84,85] and their impact on non-communicable diseases [86,87,88,89,90,91,92,93,94,95,96,97,98,99,100,101,102,103,104,105,106,107,108,109,110,111,112,113,114,115,116]. * PubMed 6, Web of Science 298, Scopus 5, Cochrane Library 6. ** Excluded by the researchers due to sample parameters (small sample) or examined/analyzed varieties that are not included in this systematic review.

3. Results

3.1. Most Cultivated Mushrooms Worldwide: Production, Alternative Source of Protein, Post-Harvest Techniques

3.1.1. The Most Commonly Cultivated Species

The most cultivated mushrooms worldwide (Figure 2) are the following [5,6]:

• Agaricus bisporus;
• Lentinula edodes or Shiitake;
• Pleurotus spp.;
• Auricularia auricula-judae;
• Volvariella volvacea;
• Flammulina velutipes;
• Tuber spp.

Fungi are a huge kingdom with over 140,000 species. The most well-known and widely consumed species are as follows: Agaricus bisporus, which includes white button mushrooms as well as brown mushrooms such as cremini and Portobello. Other notable commercial species include Lentinus edodes, commonly known as shiitake mushrooms, Volvariella volvacea (straw mushrooms), Pleurotus ostreatus (oyster mushrooms) and Flammulina velutipes (enoki mushrooms) [51]. In Greece, mushroom production is a relatively new agro-industrial sector, which emerged about 60 years ago [67]. The first cultivation techniques for mushrooms are mentioned in the 1960s for Agaricus bisporus and in the 1990s for the genus Pleurotus [67]. Mushroom cultivation is carried out in all countries of the world, with the highest production observed in China (36.2%), the United States of America (43.8%), Italy (12.8%), the Netherlands (4.2%) and Poland (3%) [51]. The cultivation of mushrooms may help to reduce our carbon footprint. Mushroom cultivation generates a large amount of CO₂ that can be used sustainably.

Mushroom production involves six sequential steps, which consist of the following: (i) phase I composting; (ii) phase II composting; (iii) spawning and spawn run; (iv) casing; (v) pinning; and (vi) cropping. Different types of mushroom require different types of substrates [25]. The description of cultivation steps is correct for Agaricus bisporus (composted substrate), but for other species, composting is not required. Depending on the variety, they are primary and secondary decomposers, and they break down organic material that plants can later use for food production. Mushrooms grow their mycelia deep into the soil, dead trees or foliage on the forest bed. The mycelia then release enzymes (proteins that accelerate chemical reactions). These special enzymes break down cellulose and hemicellulose, a building material found in the cell walls of all plants. Some fungi, such as Phellinus pini, Phlebia spp., Pleurotus spp., Klebsiella pneumoniae and Ochrobactrum tritici, can also decompose lignin, another complex structure found in the cell walls of bark and wood. This process can take anywhere between a few hours to a couple of years [5,6,7,12,13,14,15,16,17,18,19,20,21,22,23,24,25,26,27,28,29,30,31,32,33,34,35,36,37,38,39,40,41,42,43,44,45,46,47,48,49,50,51].

3.1.2. Agaricus bisporus

White button mushrooms are the most commonly consumed variety [5,26] and can grow in containers using compost as the substrate [25]. They prefer cooler temperatures [27], around 20–26 °C [31] for the incubation phase [28], and slightly warmer temperatures for fruiting [29], with 80–85% relative moisture [30]. White button mushrooms can be ready for harvest within 3–4 weeks of inoculation, making them a favorite among beginners [31]. October to March is the optimum period for growing, but this range of months refers to wild species. In controlled environments, fruiting can occur year-round [32]. However, excessive moisture can cause the substrate to become very wet, increasing mold growth [25].

3.1.3. Lentinula edodes

The best substrate is tree trunks [33], which is a common practice, but it is imperative to ensure that the wood does not contain unsafe amounts of heavy metals or pesticide residues [34]. Based on the optimal temperature range for fruiting body formation, L. edodes can be classified into different temperature types, including high-temperature type (H-type, fruiting at 15–25 °C), medium-temperature type (M-type, fruiting at 10–20 °C), low-temperature type (L-type, fruiting at 5–15 °C) and broad-temperature type (B-type, fruiting at 5 to 25 °C) [35]. Lentinula is dried to better preserve its aroma. In Japan it was the main export agroforestry product [36,39]. Today, China has taken over with the large quantities it produces [37], consumes and exports, providing a main or supplementary income to many [38].

3.1.4. Pleurotus spp.

The genus Pleurotus is found in nature with a variety of species that differ in shape, color and size. The most common are P. ostreatus, P. eryngii, P. pulmonarius, P. cornucopiae, P. columbinus, P. flabellatus, P. sajor-caju, etc. [40]. Because they are lignocellulosic decomposers, such as with broadleaf hardwood sawdust and straw-based substrates with added supplements or alder wood and beech wood [41], they are often found in tree trunks (fir), even felled trees (poplar) [42]. The Pleurotus mushroom is one of the most productive mushrooms produced by humans worldwide [43]. It may not be as productive as the white button mushroom Agaricus, however, which can produce up to seven harvests a year, compared with four for Pleurotus mushroom. However, its production is simpler, and the investment required is lower [44,45].

3.1.5. Auricularia spp.

Auricularia is an edible and medicinal mushroom in Eastern countries [12]. Its homeland is China, which produces and exports large quantities of it [13]. It grows in decaying trunks in forests, but also in courtyards of houses or covered areas [14]. Its fruit is waxy and crispy [15]. Two of the best species are cultivated: A. Polytricha and A. auricular-judea [16]. Experience shows that the second species is more desirable and is sold at higher prices than the first. A. polytricha grows at lower temperatures, 22–32 °C, and A. auricula-judae is, therefore, cultivated at a higher altitude (Figure 3). Cultivation of A. auricula-judae involves two important methods: spawn production and fruiting body production. Mycelia obtained from tissue culture were used to produce grain spawns. Environmental conditions required in intensive cultivation are also dealt with, such as the effect of temperature, humidity, light and CO₂ [17].

3.1.6. Volvariella spp.

There are 100 species all over the world of Volvariella [23]. It is considered a fungus of warm regions and grows at temperatures of 32–35 °C [23,24]. Its fruiting takes place in a short time, as it only takes two weeks from sowing to harvest, rarely longer. This is not possible with any other fungus, but with Volvariella spp. it is easy under the right conditions. It can be grown with a variety of substrates, like cotton waste, rice straw, oil palm pericarp and banana leaves [24]. However, wheat and rice straw are the best materials. For maintaining the required temperature (30–35 °C) and the fruiting temperature of nearly 28–32 °C, the main method remains the same in all cases. The morphological characteristics change according to the stage of maturity of the fungus [22]. The size of the cap varies between 6 and 12 cm, depending on nutrition and environmental conditions. The nitrogen content of the straw often varies as a percentage of dry matter, expressed as a percentage (%) [46]. This depends on the cultivation techniques, the environmental conditions, soil temperature, straw nitrogen content and straw C/N ratio [47]. So, when researchers mention straw or sawdust, it is completely impossible for each batch to contain the same nutritional value. This can only be obtained by the professional who produces the substrates and is able to analyze in the laboratory the nutrients (moisture, etc.) of the materials he uses, even those of the tree trunks, as well as their humidity. This does not mean that a layman should rely only on using ready-made substrates [48]. It is worth trying to make substrates yourself, and gradually you will gain the experience to distinguish the quality [49], e.g., of the straw or the moisture of the trunk, even without analysis [46].

3.1.7. Flammulina velutipes

This is a typical low-temperature mushroom that requires low temperature to induce fruiting body formation. Under good environmental conditions, its cap can reach up to 40 cm. As it matures, it acquires a yellow or brown color with shades of red. Its stem is cream-colored, thick at the base and becomes hollow as it matures. The substrate is the same as that used for Pleurotus, namely wheat straw. Its production lasts up to 4 months. During this period, from the fertilisation to the harvest, the humidity of the substrate is maintained through frequent but minimal watering. The total production reaches 15% of the weight of the substrate in fresh mushroom rooms [18].

3.1.8. Tuber spp.

Some species of exceptional culinary interest resist attempts to grow them in a nutrient substrate [19]. These are exclusively symbiotic fungi that require special conditions to bear fruit, in particular a symbiotic encounter with a coniferous or deciduous forest species [20]. Efforts to cultivate the Tuber melanosporum and related species have been relatively successful in artificial plantations and in symbiosis with various oak or hazelnut species [19,50]. Particularly promising is the attempt to cultivate the fungus Lactarius delicious in a symbiosis. To date, only a few species of Tuber ssp. and epigeous mushrooms, such as Tuber melanosporum, Lactarius deliciosus and Lyophyllum shimeji, are cultivable, whilst many others defy cultivation. The seed tubers should be planted 10–15 cm deep, with a row spacing of 20–30 cm and a row spacing of 75–90 cm. These guidelines ensure sufficient space for the tubers to expand and simplify field work. The time of harvest depends on the variety and climatic conditions, with maturity generally being reached in 90–120 days. The development of the tubers only takes place in a temperature range of 20 to 30° C; the optimal is 25° C, while below 10° C it is generally interrupted [6]. The average production of Tuber spp. is 4–5 kg per field, which can reach 10–12 kg with successful cultivation. Producer prices for Tuber spp. such as black truffles are EUR 600–700 per kilo and for summer truffles EUR 100–150 per kilo [21]. Lack of water in summer has a major impact on the production of black truffles in the Mediterranean region [50].

3.2. Most-Cultivated Mushrooms Worldwide: Alternative Source of Protein and Comparison with a Plant-Based Protein Source

Alternative Source of Protein

Cultivated mushrooms can be used as an alternative source of protein [52], but also as a nutritious food of choice due to their high micronutrient content [8]. The nutritional profiles of the seven most commonly cultivated mushrooms cover 15% of protein intake according to the RDI (

Table 3. Proximate analysis of the seven most popular mushrooms by dry weight basis percent (edible and wild in the first row and cultivated in the second row).

Mushroom Species	Ash (g/100 g)	Energy (Kcal)	Protein (g/100 g)	Sugar (%)	Carbohydrates (g/100 g)	Fat (g/100 g)	Salt (g/100 g)	References
Agaricus bisporus edible	9.3	336 Kcal/100 g— 1406.76 KJ/100 g	25.1	<0.1	52.7	1.7	<0.1	[8]
Agaricus bisporus cultivated	8.7	303–324 Kcal/100 1268.6 g— 1356.52 KJ/100 g	14.0–36.3	<0.1	50.9–74	0.8–2.5	<0.1	[24,53,60,62,76]
Lentinula edodes edible	6.1	340 Kcal/100 g— 1443 KJ/100 g	20.7	<0.1	59.5	1.3	<0.1	[8]
Lentinula edodes cultivated	6.7	386 Kcal/100 g— 1616.1 KJ/100 g	4.4–20.5	<0.1	67.9–87	1.7–6.3	<0.1	[76,77]
Pleurotus ostreatus edible	8	421 Kcal/100 g— 1762.64 KJ/100 g	13.2	<0.1	75.11	3.58	<0.1	[77]
Pleurotus ostreatus cultivated	6	416 Kcal/100 g— 1762.64 KJ/100 g	7–23.8	<0.1	51.9–85	0.5–5.4	<0.1	[76,77]
Pleurotus eryngii edible	6	421 Kcal/100 g— 1762.64 KJ/100 g	16.2–26.6	<0.1	−64.9	−3.5	<0.1	[78]
Pleurotus eryngii cultivated	6	421 Kcal/100 g— 1762.64 KJ/100 g	11–22	<0.1	70.5–81.4	1.45–1.57	<0.1	[76,77]
Pleurotus citrinopileatus edible	7.9	330 Kcal/100 g— 1395 KJ/100 g	37.6	<0.1	36.3	2.2	<0.1	[8,24]
Pleurotus citrinopileatus cultivated	7	421 Kcal/100 g— 1762.64 KJ/100 g	11–22.1	<0.1	52.7	2.3	<0.1	[77,78]
Pleurotus ferulae edible	5	330 Kcal/100 g— 1395 KJ/100 g	30.3	<0.1	47.8	5.7	<0.1	[53,77]
Auricularia auricula edible	3.6	140 Kcal/100 g— 586.15 KJ/100 g	12.5	<0.1	66.1	1.7	<0.1	[57]
Auricularia auricula cultivated	3.6	97–140 Kcal/100 g— 406.12–586.15 KJ/100 g	5.7–15.5	<0.1	77–91	0.4–4.5	<0.1	[53,54,55,56,57,61,76]
Volvariella volvacea edible	8–10	346.04 Kcal/100 g— 1448.8 KJ/100 g	28–32	<0.1	50–52	2–4	<0.1	[64]
Volvariella volvacea cultivated	8	346.04 Kcal/100 g— 1448.8 KJ/100 g	32	<0.1	56.8	5.7	<0.1	[22,23,24,62,63,64,76]
Flammulina velutipes edible	9.4	346 Kcal/100 g— 1448.63 KJ/100 g	17.89	<0.1	70.85	1.84	<0.1	[77]
Flammulina velutipes White cultivated	0.8	467 Kcal/100 g— 1372 KJ/100 g	23.8	<0.1	73.8	1.9	<0.1	[73,75,76]
Flammulina velutipes Brown cultivated	0.68	467 Kcal/100 g— 1372 KJ/100 g	24.8	<0.1	76.2	1.7	<0.1	[73,75,76]
Black Truffle edible	5	345.20 kcal/100— 1443.48 KJ/100 g	20.9–23.2	7.32–6.35	51.7–49.6	5.58–6.59	<0.1	[65,66,68,72]
Black Truffle cultivated (T.pseudohima layense)	8.77	339.6 Kcal/100— 1421.84 KJ/100 g	14.28	23.89	74.4	2.55	<0.1	[115]
White truffle edible (Tirmania nivea)	5.5	368 kcal/100— 1540.74 KJ/100 g	11.97	5	68	2.15	<0.1	[69]
White truffle cultivated (T.latisporum)	8.33	378.64 kcal/100— 1585.29 KJ/100 g	14.64	50	74.63	2.4	<0.1	[67,69,71,115]
White truffle cultivated (T. subglobosum)	8.13	378.6 kcal/100— 1585.12 KJ/100 g	10.96	29.98	78.68	2.23	<0.1	[115]

), and the 31 studies included in this systematic review have focused on their nutritional profile [8,22,23,24,52,53,54,55,56,57,58,59,60,61,62,63,64,65,66,67,68,69,70,71,72,73,74,75,76,77,78].

Almost 200 species of wild mushrooms are used as medicine due to their excellent nutritional profile [64]. There is variation according their protein content [115]. In addition, there is variation as concerns Tuber spp. Their phenolic content ranges from 735.01 mg/100 g d.w. to 1355.43 mg/100 g d.w. [115], and these non-nutritive components can reduce inflammation and, as a consequence, non-communicable diseases. In addition, they are a good source of selenium, magnesium and Vitamin D [8]. A total of 34.7% of total essential amino acids has been found in A. auricula-judae [53], but Volvariella volvacea [64,77] and Pleurotus ferulae [53,77] are the best sources, and for the other mushroom species, the total amount of essential amino acids was between 27% and 31.1% (

Table 4. Amino acid content (mg/100 g of protein) and portion of essential amino acids (% of total amino acids) in protein content of seven mushroom species.

Mushroom Species	Val	Leu	Ile	Thr	Met	Lys	Phe	Try	His	Total	References
Agaricus bisporus	3.9	6	3.1	3.4	1.4	6.1	3.1	-	4.23	27	[52,77]
white	5	6.3	3.8	4.6	1.4	5.9	4.4	-	4.12	31.4	[52,77]
brown	5	6.1	3.6	4.3	1.3	5.3	4.9	-	4.11	30.5	[52,77]
Lentinula edodes	3.8	6.4	3.3	5.6	2.2	5	3.8	1.9	4.22	32	[52,77]
Pleurotus ostreatus	4.7	6.8	4.3	5	1.9	6	4.3	1.4	4.44	34.4	[52,77]
Pleurotus ferulae	11.8	22.3	11.38	8.06	3.62	12.9	8.68	-	4.47	37.1	[53,77]
Auricularia auricula	3.53	4.89	1.89	4.89	0.29	4.04	2.76	-	2.16	34.7	[53,77]
Volvariella volvacea	3.5	7.55	3.3	4.88	1	5.2	6.22	1.7	2.16	35.8	[64,77]
Flammulina velutipes	4.6	6.1	4.4	4.7	1.4	5.7	4.7	1.5	2.3	34.8	[74,75,77]
Truffle black	4.9	6.1	3.6	4.6	1.5	5.5	4.9	-	2.17	31.1	[65,66,67,70,117]
Truffle white	4.9	6.1	3.6	4.6	1.5	5.5	4.9	-	2.17	31.1	[70]
Standard protein (FAO/WHO)	5	7	4	4	3.5	5.4	6.1	1	1	36	[118]

Val: valine; Leu: Leucine; Ile: isoleucine; Thr: threonine; Met: methionine; Lys: Lysine; Phe: phenylalanine; Try: tryptophan; His: Histidine.

) [65,66,67,70,74,75,77,117].

Of the 20 amino acids found in nature, 9 of them cannot be synthesized in the human body and must be obtained through dietary elements. Although mushrooms contain some of the essential amino acids, they fail to provide all nine. So, mushrooms do contain a significant amount of protein, but their protein profile is not considered complete, despite the fact that they are superior in other bioactive compounds (

Table 5. Comparison of dried shiitake mushroom (100 g/DM) vs. steak (100 g/cooked).

Steak [58]	Dried Shiitake Mushrooms [59]
20% Calories from proteins	20% Calories from proteins
Saturated Fats	Unsaturated Fats
Higher lipidemic content	Lower lipidemic content
No Fiber	High in fiber
Iron 0.79 mg	Iron 18 mg
Magnesium 17 mg	Magnesium 17 mg
Calcium 35 mg	Calcium 22 mg
Potassium 288 mg	Potassium 464 mg
Vitamin C 0 mg	Vitamin C 25 mg
Depletes the earth	Adds needed nitrogen to the soil

) [8]. Special care is required by vegetarians. They should combine different foods (lentils with rice, corn with grains, mushrooms with broccoli and corn, etc.) to meet 100% of their protein needs [52].

Nowadays, cultivated mushrooms are popular and valuable foods because they are low in calories, carbohydrates, fat and sodium. Mushrooms also provide important nutrients, including selenium, potassium, riboflavin, niacin, vitamin D, proteins and dietary fibers (chitin, chitosan, glucans) and other bio-active compounds with medical value [114]. All the cultivated mushrooms mentioned in this systematic review have high amounts of dietary fiber. Dietary fiber-rich foods associated with physiological actions in the small and large intestine have important implications for human health. These properties include water dispensability and solubility, viscosity effects, bulk, absorption, ferment ability and binding of other harmful compounds such as cholesterol, sugars and fat. These properties may lead to various physiological actions, such as reducing cholesterol and alleviating blood glucose elevation, maintaining gastrointestinal health and positively affecting calcium bioavailability and immune function [119].

On the other hand, mushrooms have been highly studied for their therapeutic properties and are a major source of income for many people, especially in Asia (Figure 2). Another form that could be utilized is as a protein powder [3]. Protein powders are particularly useful for active people or those with higher protein requirements (e.g., athletes, older adults) who find it difficult to meet their needs through whole foods, but they are the only source of high protein content for vegetarians, and compared with some other sources like beans, they do not have have side effects [120]. A study by Morton et al. (2018) highlights the effectiveness of protein supplementation in promoting muscle protein synthesis, particularly during exercise [121]. For those with busy schedules, protein powders offer a quick and easy way to prevent muscle breakdown and stay energized. For vegans, vegetarians or those with limited dietary options, protein powders can be a valuable source of complete protein. For most people, protein powders are a convenient supplement, but not a necessity. Whole, nutrient-dense foods should remain the primary source of protein [120].

3.3. Most-Cultivated Mushrooms Worldwide: Post-Harvest Techniques with Important Benefits and Low Cost

Post-Harvest Techniques to Improve Their Nutritional Profile

Ten studies included in this systematic review have shown that cultivated mushrooms are rich in vitamin D with proper post-harvest practices (

Table 6. Cultivated mushrooms and their content of Vitamin D2 (µg/100 g fresh weight), ergo sterol, ergo sterol ergosta7,22-dienol, ergosta-5,7dienol, ergosta-7enol (mg/100 g f.w.) [62,80,81,83].

Mushrooms Species	Vitamin D2	Ergo Sterol	Ergo Sterol Ergosta7,22-Dienol	Ergosta-5,7dienol	Ergosta-7enol
Agaricus bisporus	0.11	56.3	1.78	6.03	1.34
Agaricus bisporus irradiated with UVB	11.2	11.2	1.73	4.7	1.28
Lentinula edodes	0.44	84.9	2.26	6.51	5.03
Pleurotus ostreatus	0.72	68.0	<1.66	8.89	<1.7
Volvariella volvacea	0.50	84.9	2.26	6.51	5.13
Volvariella volvacea irradiated with UVB	19.2	19.2	1.73	4.7	1.28
Flammulina veluptipes	0.14	35.5	<1.49	16.5	2.32

and Figure 4) [22,24,62,79,80,81,82,83,84,85]. Vitamin D, often referred to as the “sunshine vitamin”, is associated with mental health. Emerging research suggests that maintaining adequate vitamin D levels may play a role in preventing the onset of depression, as it influences the production of neurotransmitters such as serotonin, which are crucial for mood regulation but also have anti-inflammatory properties [122]. Vitamin D is also a hormone [123] that helps control cells and organ function [124]. Ergocalciferol (vitamin D2) is the form of vitamin D that can be formed by exposure to ultraviolet radiation [125]. New vitamin D derivatives can be formed by different metabolic processes and an appropriate diet could help to prevent deficiency [122]. On the other hand, there are many other techniques, for example, cooling of mushrooms, freezing of mushrooms, edible coatings of mushrooms, washing of mushrooms with antimicrobial agents, ozone and electrolyzed water, packing of mushrooms, irradiation, etc., for preserving and improving the nutritional profile of mushrooms [123].

Another post-harvest technique is drying mushrooms, and the best drying temperature to preserve their nutritional value is 60 °C, as the protein and carbohydrate content is well preserved [22,23,24].

Food intolerance is increasingly talked about, but self-diagnosis can lead to unnecessary dietary restrictions, nutritional imbalances, and even worsening health [8]. Eliminating major food groups like dairy without proper substitution, like a combination of mushrooms [79], can lead to lack of essential nutrients, such as calcium, which is necessary for vitamin D absorption [124]. Additionally, an inadequate intake of vitamin D is usual, especially in winter [124], and sundried mushrooms are an easy solution [79], as shown in

Table 7. Main food sources of vitamin D [124].

Food Sources of Vitamin D	Vitamin D (μg/100 g)
Fish
Cod liver oil	210–250
Salmon	13.1–24.7
Farm salmon	6.0
Herring	5.7–15.4
Smoked salmon	5–27
Dairy products
Full-fat milk	0.1
Butter	3.4–8.4
Egg yolk	0.5–5.4
Beef liver	1.2
Mushrooms
Agaricus bisporus fresh	0.7–2.3
Lentinula edodes sundried	40

.

In such cases, an enriched shiitake powder was the solution [79].

3.4. Therapeutic Properties of the Most-Cultivated Mushrooms Worldwide and Comparison with Wild Mushrooms as Concerns Their Pharmaceutical Value

3.4.1. Non-Communicable Diseases

Mushrooms’ impact on non-communicable diseases is valuable and obvious from the findings of the thirty-one studies included in this systematic review (

Table 8. Mushrooms’ impact on non-communicable diseases (both in vivo and in vitro studies).

Authors (Year)	Mushroom Species	Medical Value	References
Khursheed et al. (2020)	Agaricus bisporus, Pleurotus ostreatus, Lentinula edodes	Improves serum/plasma triglycerides and hs-CRP.	[93]
Ahmad et al. (2023)	Lentinula edodes	Acts as an antimicrobial, antiviral, anticancer/antitumor, antidiabetic, antihyperlipidemic, anticholesterol	[87]
Garcia et al. (2021)	Lentinula edodes	May be potential source of natural antioxidants, antibacterial agents and anti-aging agents.	[88]
Zeng et al. (2012)	Pleurotus eryngii, and Flammulina velutipes	The methanolic extracts of the dried caps of the mushrooms were determined using a number of different chemical reactions in evaluating multi-mechanistic antioxidant activities	[89]
Ziaja-Sołtys et al. (2020)	Lentinula edodes	Anticancer properties from Lentinus edodes fruiting bodies on human breast cancer	[90]
Łysakowska et al. (2023)	Shiitake (Lentinula edodes)	Acts as anti-obesogenic, antiviral, anticancer/antitumor, antidiabetic, antihyperlipidemic, anticholesterol	[91]
Martel et al. (2017)	Agaricus bisporus, Pleurotus ostreatus, Lentinula edodes	Anti-obesogenic and antidiabetic effects	[92]
Khursheed et al. (2020)	Pleurotus spp.	The polysaccharides from mushrooms are effective against type ΙΙ diabetes mellitus by reducing oxidative stress and also act as prebiotics and reduce gut dysbiosis, thereby helping in managing insulin resistance and type ΙΙ diabetes mellitus	[93]
Elkhateeb et al. (2023)	A. auricula-judae, L. edodes	Antidiabetic properties of mushroom species were promising	[94]
Khan et al. (2023)	Auricularia cornea	The crude polysaccharides from A. cornea mushroom strains act as natural antioxidants	[95]
Anwar et al. (2019)	Lentinula edodes	Total cholesterol and LDL cholesterol concentrations were reduced	[96]
Yadav et al. (2020)	Pleurotus giganteus	Protective effect against neuronal dysfunction	[97]
Kundu et al. (2021)	Agaricus bisporus, Pleurotus spp.	Source of important nutrients having hepatoprotective and antihyperlipidemic actions	[98]
Lee et al. (2019)	Pleurotus giganteus	Decrease neurotoxicity through various neuroprotective molecular mechanisms	[99]
Ye et al. (2024)	Flammulina velutipes	Immunomodulatory, anti-inflammatory and antibacterial properties	[100]
Yamac et al. (2010)	Agaricus bisporus	Pancreas protective effect of button mushroom Agaricus bisporus	[101]
Chugh et al. (2022)	Agaricus arvensis and Agaricus silvaticus	130 medicinal activities like antitumor, immunomodulation, antioxidant, radical scavenging, cardioprotective and antiviral actions	[102]
Won et al. (2029)	Shiitake	Protection against osteoporosis	[103]
Lindequist et al. (2021)	Lentinula and Pleurotus	May be able to improve bone stability by influencing different steps of bone formation, mineralization or resorption.	[104]
Qu et al. (2019)	Auricularia auricula	May be protection against osteoporosis	[105]
González-Ibáñez et al. (2023)	Pleurotus ostreatus	Pleurotus ostreatus reduces endoplasmic reticulum stress and inflammation in adipose tissue of obese subjects	[106]
Merdivan et al. (2017)	Agaricus subrufescens, Flammulina velutipes, Pleurotus ostreatus and P. pulmonarius	Antiallergic activities	[107]
Seo et al. (2021)	Lentinus (Lentinula), Auricularia, Flammulina, Pleurotus, Agaricus	Some mushroom compounds that act against HIV, influenza A virus and hepatitis C virus showed antiviral effects comparable to those of antiviral drugs	[108]
Yan et al. (2017)	Tuber latisporum, T. subglobosum and T. pseudohimalayense	Antioxidant properties	[115]
Rauf et al. (2023)	Agaricus bisporus, Pleurotus ostreatus, Lentinula edodes, Flammulina velutipes	Helpful in alleviating hypertension and other cardiovascular malfunctions	[114]
Blumfield et al. (2017)	Agaricus bisporus	Beneficial impact on biomarkers correlated with metabolic syndrome and gastrointestinal health	[111]
Atila et al. (2017)	Agaricus bisporus	Has been reported to have antimicrobial, anticancer, antidiabetic, antihypercholesterolemic, antihypertensive, hepatoprotective and antioxidant activities	[112]
Wong et al. (2020)	Agaricus bisporus, Pleurotus abalonus, Pleurotus eryngii, P. ostreatus	Antineoplastic effectiveness in human clinical trials	[113]
Dimopoulou et al. (2022)	Agaricus bisporus, Pleurotus citrinopileatus, Lentinula edodes	Act as an antimicrobial, antiviral, anticancer/antitumor, antidiabetic, antihyperlipidemic, anticholesterol	[8]

), without ignoring the side effects of overconsumption (over 1 Kg per day) [86,87,88,89,90,91,92,93,94,95,96,97,98,99,100,101,102,103,104,105,106,107,108,109,110,111,112,113,114,115,116].

The medicinal values in Table 8 relate only to nutritional components or to non-nutritive components such as phenols, terpenes, sterols, non-protein amino acids, nucleosides, etc., due to the fact that only a few studies [2,116] took place with patients as participants.

3.4.2. Cardiovascular Disease

The number one cause of death worldwide is cardiovascular diseases, which are related to lifestyle habits (diet, smoking, low physical activity) [86]. In recent studies, Lentinula edodes was found to reduce the risk of cardiovascular diseases if it is consumed daily as a plant-based source of protein [87].

Recent guidelines state the importance of replacing saturated fats with monounsaturated and polyunsaturated fats for primary and secondary cardiovascular prevention. Although studies are still needed to confirm the data, it certainly seems to be mainly due to the low caloric content of mushrooms, the high content of micronutrients and their high antioxidant activity that they are suggested for the prevention of cardiovascular disease. Their antioxidant action seems to be due, on the one hand, to the antioxidant vitamins and, on the other hand, to phytochemical substances [88]. Most of the cultivated mushrooms contain 1.5–2.4 g/100 gr d.w. when they are fresh [8]. The recommendations state that a diet containing about 25–30 g dietary fiber per day, from various sources such as mushrooms, can significantly reduce the risk of cardiovascular disease. The studied mechanisms of action concern weight reduction, inflammation and oxidation, the effect on glucose metabolism and the lipid profile [86].

Moreover, the high concentration of potassium that A. bisporus, A. blazei and Pleurotus ostreatus contain reduces blood pressure and has cardio-protective effects. A medium-sized mushroom (90 g) contains more potassium than a banana or a glass of orange juice. It also contains copper, which participates as a shield for many functions of the heart muscle [89]. The doses that could have cardio-protective effects range for mushrooms Ganoderma lucidum, Agaricus brasiliensis and Cordyceps sinensis from a 400 μg/mL dose to a 200 μg/mL dose, or even 10 μg/mL, respectively [116].

3.4.3. Cancer

Cancer is one of the most important causes of death worldwide and the second highest cause of death in Greece, after cardiovascular diseases [1]. Although mushroom consumption appears to play a role in some forms of cancer, such as colon and stomach cancers, the etiology of carcinogenesis is multifactorial and is influenced by a combination of a multitude of environmental, metabolic and genetic factors [90]. Mushroom extracts have been studied as influencing factors in the pathophysiology of carcinogenesis at all stages, possibly affecting carcinogen metabolism, cell division and tumorigenesis. Mushrooms contain substances that can resurrect carcinogenesis (antioxidants, vitamins, minerals, physicochemicals) [8]. Compared to the increased consumption of red meat, which has been associated with a possible occurrence of colon cancer, mushrooms, which are now considered a vegetable source of protein, seem to protect against this type of cancer because they are rich in vegetable fiber [87].

Also, the antioxidants of mushrooms, such as vitamins A, C, E and β-carotene, seem to have a role in inhibiting the pathophysiological mechanisms of cancer, although more studies are needed for safer conclusions [91]. The intake of 100 g per day of dried mushrooms appears to lead to a lower risk of developing cancer based on their nutritional assessment [8].

Shiitake and Reishi mushrooms in particular probably have anticancer properties. Many studies have shown that a component of shiitake mushrooms, lentinan, can stimulate the immune system to fight viral infections and cancer cells. In fact, lentinan is already used as an anticancer drug in Japan [90]. The first research to show that mushrooms have anticancer activity was published in the 1960s [87]. Long-term research has identified some of the compounds responsible for this activity, such as polymers, lectins and terpenes. The conclusions from this research suggest that the probability of developing breast cancer decreases with the consumption of mushroom fruiting bodies [90].

The most recent data report the protective effect of mushroom consumption on various forms of cancer, such as stomach, ovarian, lung, prostate and liver. Their antitumor activity, mainly in human cell lines, is due to the polysaccharides found in a variety of mushrooms [8]. These polysaccharides are in the form of β-glucans and have been shown in clinical trials to improve cellular immunity through the activation of T-helper cells and the production of interferon-γ and interleukins. They also stimulate the activity of killer NK cells and the gene synthesis of the enzyme nitric oxide synthase [87]. Only 6 and 9 g/day of C. versicolor mycelium powder had anticancer effects [102]. The anticancer activity dose was different for different mushroom species; specifically, for Pleurotus ostreatus it was 500 µg/mL [116].

3.4.4. Diabetes Mellitus

Diabetes mellitus (DM) has become one of the most important causes of premature mortality in most countries, mainly due to an increase in the risk of developing cardiovascular disease [92]. Lack of information about DM, combined with inadequate nutrition education, can lead to complications such as blindness, amputation and kidney failure [93]. A growing body of evidence shows that in people with pre-diabetes, eating mushrooms can delay or help prevents the onset of type 2 diabetes mellitus [94]. Antidiabetic properties were investigated for mushroom species A. blazei Murrill and Auricularia spp., with positive effects on glucose control [94,95]. The highest effective daily intake of mushroom powder was 3 × 2.215 g/day and could be compared with Acarbose [94].

Mushrooms have been investigated because of their low saturated fat and low glycemic index. Their glycemic index is close to zero [95]. The amino acids included in mushrooms have also been studied, with the aim of investigating the positive benefits in blood glucose control and insulin sensitivity. The chromium they contain may be effective in optimizing insulin metabolism and reducing plasma cholesterol levels [92].

Specific polysaccharides produced by mushrooms have antidiabetic effects, since they reduce plasma glucose and enhance insulin sensitivity [8]. In addition, they reduce triglycerides, LDL and total cholesterol in diabetic clinical animal models, while increasing HDL cholesterol [93]. The dose with antidiabetic effects for Agaricus blazei mushroom extract is 1500 mg/d [102]. Finally, antidiabetic effects succeded with a dose of 100 µg/mL Ganoderma lucidum extract or 50 µg/mL Inonotus obliquus extract or 400 µg/mL Pleurotus florida extract [116].

3.4.5. Dyslipidemia

One of the reasons for the development of cardiovascular diseases is LDL fraction and high levels of “bad’’ cholesterol, and mushrooms contain substances that can lower them [8]. These include both high-molecular components of the cell walls (glucans, glycans, chitin or chitosan) and low-molecular secondary metabolites [96]. Eritadenine (alkaloids) produced by the shiitake mushroom accelerates cholesterol metabolism in the liver, which in turn leads to a reduction in its quantity in the blood plasma [126]. In addition to their numerous beneficial properties, they are also characterized by a low calorie and fat content. They are classified as foods with a low total cholesterol content and are, therefore, suitable for consumption by patients with dyslipidemia. Lentinus edodes, Pleurotus spp., Coriolus spp. And Schizophyllum spp. Are good examples of medicinal mushrooms. The addition of mushroom powder (500 g/kg/body weight, but dose depends on patient age and other medical conditions) to a hypercholesterolemic diet had health benefits [98]. In another study, 50 g of cooked Pleurotus ostreatus three times regularly over 24 days was sufficient to lower cholesterol, blood sugar and triglycerides in diabetics [102].

3.4.6. Hypertension

Research from Harvard University has shown that mushrooms are a very tasty food, with no added salt and minimal sodium content [8]. By frequently adding them to foods instead of salt, the total sodium intake during the day can, therefore, be reduced [114]. The dose of mushroom extract that can act as a natural antioxidant ranges from 1 mg to 5 mg, or 6 GAE for the mushrooms Inonotus obliquus, G. lucidum and A. bisporus [116]. Most importantly, novel foods were used in clinical trials, such as milk powder derived from Pleurotus ostreatus, and not only supplements produced from G. frondosa, P. eryngii and H. erinaceus. Biomarkers used to determine the causal relationship between diet and hypertension are cholesterol, total LDL, HDL, fasting triacylglycerol, homocysteine, homeostasis and platelet aggregation [114].

3.4.7. Neurological Disorders

(a)

Scientific studies have shown that the human brain and the cells of the nervous system are particularly sensitive to oxidative stress because of the following [97]:

(b)

The brain accounts for 2% of total body weight and consumes 20% of the oxygen supply.

(c)

It is rich in polyunsaturated fatty acids, which are easily oxidized.

(d)

It does not have a strong antioxidant defense in its cells.

(e)

It contains large amounts of iron and ascorbate, which catalyze lipid peroxidation.

(f)

Many neurotransmitters auto-oxidize and produce free radicals.

(g)

In some areas, nerve cells are damaged by mitosis and oxidative damage.

Through the activation of genes, such as those associated with the production of NO synthase, interleukin IL-1b and tumor necrosis factor (TNF-a and NF-kB), an inflammatory process takes place in the nerve cells, leading to the production of radicals and cytokines. Through this process, mushrooms and nerve cells appear to influence the oxidative process and reduce the risk of diseases such as Alzheimer’s and Parkinson’s [99]. On the other hand, the immunomodulatory potential could also have a positive effect on general health, and the dose of each mushroom ranges from 20 µg for the extract of Pleurotus ostreatus to 25 µg/mL for the extract of Grifola frondosa and 30 µg/mL for the extract of Lactarius deliciosus [116].

Another and more important property of mushrooms is their use as a narcotic and hallucinogenic material. Only a few years ago, the mushroom A. bisporus was considered unique in this respect [126], as was F. velutipes [100]. Today, medicinal mushrooms such as Pleurotus giganteus and Hericium erinaceus are produced worldwide as dietary supplements for neurological disorders [97].

In recent years, however, a number of other psychotropic mushrooms belonging to the genera Psilocybe, Conocybe, Panaeolous, Strophalia and Mycena have been intensively studied, and various medicinal substances such as psilocybin and psilocin, which are currently used for therapeutic purposes in neuropsychiatry, have been isolated from edible mushrooms [102]. Thirty women who consumed four cookies containing 0.5 g of H. erinaceus powder daily for one month were found to have lower levels of anxiety, panic and depression, but Agaricus blazei also had a positive effect on anxiety and mental stability [102]. In another study with 1800 mg three times a day for another mushroom (G. lucidum), a reduction in the Clinical Global Impression (CGI) severity scale was found [99]. Finally, extracts of Hericium erinaceus (100 µg/mL), Phellinus rimosus (50 µg/mL) and Cantharellus cibarius (100 µg/mL) had a neuroprotective effect [116].

3.4.8. Osteoporosis

Numerous studies have shown that women can prevent osteoporosis if they increase their vitamin D and calcium intake [103]. It has been known for some time that bone loss is greater in the winter months for two reasons. Firstly, our bodies are exposed to less sunlight, which means that the body does not produce the same amount of vitamin D. Secondly, our bodies do less exercise in winter months, which also reduces bone mass. Researchers wanted to find out whether “winter bone loss” can be treated therapeutically with a diet [104].

In their study, the subjects were given calcium supplements made from mushrooms, which increased their total daily calcium intake to 800 g. Half of the subjects in the group received a vitamin D supplement containing 400 IU, while the other half received an inactive drug. After six months, the bone and spine density of both groups was measured, and the results showed that the women who took a vitamin D supplement during the winter months had lost only half as much bone mass as those who took only calcium [105].

The vitamin D2 content in mushrooms [80] ranges from 0.11 μg/100 g f.w. for Agaricus bisporus to 0.72 μg/100 g d.w. for Pleurotus ostreatus. Exposing the mushrooms to sunlight during the drying process increases the vitamin D content of the mushrooms, an essential component for the proper functioning of the immune system, but also for osteoporosis [103].

3.4.9. Other Effects

The consumption of mushrooms stimulates the function of the T-lymphocytes and killer cells of our immune system when there is a defense situation [100]. On the contrary, in mild situations where the organism is safe, it does not attack unnecessarily and does not cause inflammation, mutations or alterations [106].

The antimicrobial properties of mushrooms promote the body’s defense against the host by enhancing cytokine secretion [100]. Antiviral properties of certain mushrooms, such as A. blazei and A. brassiliensis, have been found in in vitro studies. Some other fungal species may not kill the virus, but act in the early stages of its replication [107]. An anti-inflammatory effect was mentioned for Cordyceps militaris at a dose of 300 µg/mL and Hericium erinaceus at 0.5 mg/L, but also for Phellinus igniarius at a dose of 100 μmol/L [116].

Scientists report that fungal lectins regulate the production of immune proteins. It is known that lectins have stable properties and are not altered by the processing of mushrooms [108]. Under the name “Agaricus muscarius”, an extract of fresh mushrooms is administered in infinitesimal doses to treat rheumatism and eye irritation [126].

It is worth noting the report that eating mushrooms reduces the development of obesity, as mushrooms are low-calorie foods that promote satiety due to their high fiber content. Mushrooms consist of 80–90% water and 8–10% fiber [8]. So, the best way to curb your appetite is to include these miraculous mushrooms in your diet.

3.4.10. Side Effects

Finally, they are rich in phosphorus and zinc compared to other vegetables. There are few data reporting clinical problems of mushroom consumption in the areas of blood, kidneys, glucose metabolism, lipids or liver. Allergic problems have been reported from mushroom seeds, but only from mushroom traders and only in Japan [109].

Mushroom poisoning can develop into an extremely serious and life-threatening condition for which there is no antidote. Depending on the symptoms of the poisoning and the damage to organs and systems, the following syndromes are distinguished [110]:

• Gastroenteritis
• Phalloid poisoning
• Muscarinic poisoning
• Mycoatropin poisoning
• Orleans poisoning
• Gyrometry poisoning
• Psilocybin poisoning

Several mushroom toxins have been identified in recent years, but these require a very high consumption of mushrooms or the consumption of wild mushrooms [110]. This mechanism of toxicity could be related to reduced glutathione and ascorbate levels, which are important protectors against oxidative damage, and Figure 5 partially explains the mechanism of this toxicity [110]. The only safe way for prevention is the collection of wild mushrooms by a professional mycologist with experience in this activity or the purchase of cultivated mushrooms with beneficial effects [111] and without side effects, if the consumption does not exceed 1 kg per day [102,112]. However, a solution is presented in Figure 5 [102].

4. Discussion

This systematic review aims to provide a brief overview of the most commonly cultivated mushrooms species in Greece and other countries (A. bisporus, L. edodes, P. citrinopileatus, A. auricula-judae, V. volvacea, F. velutipes, Tuber spp.) in terms of their nutritional composition as well as their medicinal and biological properties (Figure 2) [127]. Production in industrialized countries is declining or has reached a plateau. Shiitake, Pleurotus species, etc., which are mainly cultivated in Asian countries, are now also cultivated in Europe [127], and the consumer demand for Lentinula edodes and Pleurotus is 10–12 tons per year, but the cultivation rate covers only 30% of the demand. Another important point is that other species such as Tuber have been cultivated and accepted due to the low cultivation costs and ease of cultivation [67].

The mushroom has managed to be one of the most delicious foods from the time of hunter–gatherers to the modern world. In addition to wild mushrooms, there are also those cultivated in intensive or extensive production units that offer everyone the opportunity to obtain what is rightly called “food medicine” [91]. They enrich the body with vitamins of high biological value and amino acids (Table 4) that protect against cholesterol, anemia, sugar, arteriosclerosis, heart muscle abnormalities and cancer [111]. They are also considered a dietary food, as one kilogram of mushrooms contains only 190 calories, with the potential to replace other fattening foods while providing a feeling of satiety [112]. All mushroom ingredients remain intact after cooking. Moreover, frying increased the protein (2.01% ± 0.2% [fresh mushroom] to 2.23% ± 0.09%), lipid (14.68% ± 0.9% to 15.56% ± 0.34%) and carbohydrate (0.89% ± 0.01% to 2.69% ± 0.03%) content, while microwaving increased the protein (2.01% ± 0.2% to 3.64% ± 0.08%) and carbohydrate content. Drying resulted in a significant increase (p < 0.05) in protein (2.01 ± 0.2% to 24.36 ± 0.09%), carbohydrate (0.89% ± 0.01% to 58.67% ± 3.29%) and phenolic contents (8.77 ± 0.1 to 119.8 ± 0.7 mg GAE/g mushroom), while freezing only increased the carbohydrate content (0.89% ± 0.01% to 1.77% ± 0.03%). Finally, boiling resulted in the lowest true retention levels of vitamins and bioactive compounds, while roasting resulted in higher true retention values. In general, the true retentions of minerals were lowest in each type of cooking method. However, the antioxidant activities of microwaved and roasted samples were maintained or increased [128]. On the other hand, there are many other techniques, for example, cooling of mushrooms, freezing of mushrooms, edible coatings of mushrooms, washing of mushrooms with antimicrobial agents, ozone and electrolyzed water, packing of mushrooms, irradiation, etc., for preserving and improving the nutritional profile of mushrooms [123].

Consumer preferences for mushrooms are based on their type and freshness. They must have a good color, they must not be completely open, they must not have passed the ripening stage, because then their skin becomes hard [128]. All mushrooms have their own characteristic aroma [129]. For spectacular culinary results from local mushrooms, Pleurotus spp. are the best raw products for salads [130]. A. bisporus and L. edodes are ideal for soups and black truffle as a garnish for fish and/or pasta. All mushrooms are suitable as ingredients in many recipes; the best result is a combination of two or three different mushroom species [129].

Although taste and physicochemical properties play a role for consumers, the second and most important criterion is their nutritional value and health benefits. This systematic review of 115 studies published from 2010 to January 2025 (Figure 1, Table 1 and Table 2) focused on fresh mushrooms that increase vitamin D content when exposed to sunlight [62,82,83,84] (Figure 3) or UV radiation from lamps (Table 6), or even dried mushrooms exposed to UV radiation from lamps [62,80,81,83], in addition to their cultivation and nutritional profile. In summary, it depends on the type and orientation of the mushrooms, whether they are sliced or whole, the distance from the lamp housing, the size of the mushroom and the total number of pulses received [83]. Previous studies have again shown that a temperature of 60 °C is suitable for drying mushrooms, as this preserves most of the nutritional value [23,24].

Nearly 200 wild mushroom species are used as medicine due to their superior nutritional profile and bioactive compounds [64], but cultivated mushrooms are easier to find and in many countries mushroom picking is prohibited by law; cultivated species might also provide health benefits. This systematic review, therefore, also examined the effects of mushrooms on NCDs and found that eating mushrooms with moderate regularity indicates potential benefits, such as reduced mortality from cardiovascular disease. Weak and very weak evidence suggests potential benefits in relation to blood pressure and blood lipids, and the long-term effects of these effects on well-being have been demonstrated (Table 8). Mushrooms as a dietary supplement may have the following health benefits [91]:

• Regulation of cholesterol and sugar levels;
• Strengthening of the immune system;
• Support for anemia;
• Prevention of various forms of cancer;
• Control of body weight;
• Protection against inflammation;
• Cartilage formation and bone calcification;
• Prevention of cardiovascular diseases;
• Slowing down the natural aging process of cells;
• Regulating the metabolism of carbohydrates, lipids and proteins;
• Good functioning of the gastrointestinal tract.

They are considered an excellent food because they contain only small amounts of fat and sugar, but considerable amounts of fiber [131]. Mushroom proteins are of high biological value due to the presence of all the necessary amino acids and rival animal proteins [52], but without the toxins, fats and cholesterol they contain, which is why they are considered ideal for vegetarians [60]. Due to their bioactive compounds, mushrooms exhibit a variety of health-promoting properties, including antioxidant, anticancer, immunomodulatory, antidiabetic, neuroprotective, antihypertensive, hepatoprotective, antifungal, antimicrobial, antiviral and antibacterial properties [113]. They are also an ideal choice for those who want to lose weight, as they contain few calories [8]. The low sodium content makes them particularly interesting for those on a low-salt diet. The lower calorie content is due to the fact that they are 95% water, while they are low in fat and carbohydrates [132].

Mushrooms also contain nutrients that make them an ideal choice for a weekly diet. Beef and pork are rich in B vitamins, more specifically vitamins B1 (mainly pork), B6, B3 and B12. Research has shown that a serving of pork contains on average 37% of the DRIs for vitamins B1, B6 and B12. Pork is one of the richest sources of vitamin B1 (thiamine), as it contains about 5–10 times the amount of thiamine found in beef [58]. Pork or beef steak may be the lowest in vitamins A, B6 and B12 [133,134], but the mushroom L. edodes and other species such as A. bisporus, L. edodes and Pleurotus ostreatus are superior to steak in vitamin C, vitamin B2, vitamin B9 and vitamin B3 [134,135]. The mushroom variety L. edodes has a higher content of iron, zinc and copper compared to 100 g of steak (Table 5) [59]. It also appears to contain as much or slightly more selenium than pork chops. Selenium prevents the occurrence of various types of cancer, has anti-aging properties, helps the thyroid gland to function properly and stimulates the immune system [136]. It contains several B-complex vitamins such as riboflavin and niacin, which are essential for the functioning of the nervous system and skin health, but also vitamin D (Table 7), which is important for bone structure [137]. In addition, mushrooms contain trace elements such as copper for metabolism and selenium to protect against oxidative stress [113]. Mushrooms either as food supplements (tablets, capsules, powdered or in extracts) [129] or as foods, per the seven cultivated mushrooms, have beneficial effects [138].

In addition to their high dietary value, the production of mushrooms can, therefore, easily mean an additional income for farming families. Considering that at a price of EUR 2–2.5 per kilogram, a product with a high protein content (over 27% of dry weight) and amino acids very close to those of meat is produced, mushrooms are an excellent means of nutrition at low cost with the proper techniques of cultivation (Figure 4) and no danger of poisoning if they are cultivated (Figure 5). The forms in which someone can utilize their production are fresh and dry. Normally the first production is 50% of the total production. After 10–14 days, the 2nd cycle of coke production begins. Within 10–12 weeks we have consumed most of our production [47] and can then switch crops. Thus, the small area of one square meter will have produced 125 kilos of mushrooms within 3 months. When you consider that the moisture content of fresh mushrooms is between 82–92% of the green weight and that of dried mushrooms is 12–13% and, therefore, the ratio of dry to fresh mushrooms together with the other relative wastes is about 1:6, it becomes clear why the mushroom trade allows for huge profit margins [139].

This systematic review underscores the importance of validating the nutritional and sustainable role of alternative protein sources (Table 3) such as cultivated mushrooms in the food supply and highlights clinical approaches to maximize their transformative impact in healthcare. Future research directions could also emphasize their sustainability over other post-harvest methods to increase their cultivation, and more clinical interventions could reveal new medical aspects of mushrooms use.

5. Conclusions

Conclusively, an appropriate diet includes mushrooms that are a rich source of protein of non-animal origin and fibers and other bioactive compounds, but also with low cholesterol content that can contribute significantly to both primary and secondary prevention of a variety of degenerative diseases. Although the scientific evidence shows that health effects are due to the totality of a diet and the interaction of foods and their components, it is clear that mushrooms play a potential role in important functional goals of the organism. As it is often difficult to fully comply with dietary recommendations for the prevention of various diseases, mushrooms can be beneficial in the prevention of diabetes, cardiovascular disease and metabolic syndrome, but always as part of a balanced dietary model. Because, as has been known since Hippocrates, nutrition is the greatest driver of imbalances in biological networks and the greatest lever for rapid change, reversing disease and creating health. New post-harvest techniques such as drying, freezing and cooking or even storage (12 °C/12 days) guarantee the extraction of higher bioactive compounds from cultivated mushrooms and thus a higher medicinal value with low cost, and their production could be a future sustainable food supply route with a low environmental footprint. That is the reason they are recognized as the new superfood for the future.