A mushroom is a spore-bearing fruiting body of a fungus—a simple definition for a rather complex group of living organisms. Indeed, mushrooms and other fungi are considered neither plant nor animal—they comprise their very own self-named kingdom within the Eurkarya domain of the taxonomy of living things.1 In fact, recent studies indicate that fungi are actually more closely related to animals than to plants and play an essential role across the various ecosystems of the world.2 Unfortunately, describing what makes the over six million species of fungi worldwide different from plants and similar to animals, as well as how they impact nature as a whole, is well-beyond the scope of this article. So, you’ll just have to settle for this review article in which we focus on edible species of mushrooms and the roles they play in human diet and health.
History of Mushroom Cultivation
Cultivation of mushrooms likely began with the Chinese—Tao Hongjing (456–536 CE) included a commentary on cultivating the mushroom Wolfiporia cocos in Bencao Jing Jinzhu, an extension of an extension of the Shennong Bencao Jing, the oldest surviving Chinese materia medica, which categorized medicinal uses of 365 herbs.3 In contrast, early cultivation of mushrooms in Europe was documented much later, during the mid-1600s, beginning in abandoned quarries and caves near Paris.4 Both Asian and European mushroom cultivation practices were subsequently introduced in the United States in the 1870s.5 Today, of about 2,000 species of mushrooms considered safe for consumption, 25 to 35 are widely eaten, and fewer than that are commercially cultivated.6,7
Mushroom Nutrient Composition
While the primary mass of most mushroom fruitbodies is water, depending on the species—and even within the same genus—mushrooms contain varying percentages of carbohydrates (3–42% dry matter [DM]), protein (4–44%; maximum, 57.3% DM), and lipids (2–6% DM).8 Mushrooms are low in calories yet contain a variety of vitamins, polyphenols, carotenoids, macroelements, and other bioactive components.8,9 In an analysis, the addition of an 84g serving of commonly consumed raw mushrooms (e.g., the white, brown/cremini, and portabella states of Agaricus bisporus) to United States (US) Department of Agriculture Food Patterns resulted in a 2- to 3-percent increase in fiber, an 8- to 12-percent increase in potassium, a 12- to 18-percent increase in riboflavin, an 11- to 26-percent increase in niacin, an 11- to 23-percent increase in selenium, and a 16- to 26-percent increase in copper depending upon the pattern type and calorie level but only one-percent or less increase in sodium, a one-percent increase in calories, and no effect on saturated fat or cholesterol.10 Adding a serving of raw specialty mushrooms (Pleurotus ostreatus) also increased dietary vitamin D by 8 to 11 percent and dietary choline by 10 to 16 percent.10
The nutrient profiles of mushrooms can also vary depending on the environment in which they grow; for example, a comparison of wild and commercial species of mushrooms revealed that the latter generally contained more fat, less protein, and more sugar. When considering types of vitamin E, higher levels of α-tocopherol but undetectable levels of γ-tocopherol were found in the wild species, and the wild species also contained lower concentrations of monounsaturated fatty acids but higher concentrations of polyunsaturated fatty acids as well as higher concentrations of phenols but a lower concentration of ascorbic acid compared to commercial mushrooms.11 A separate study confirmed the greater phenol content and antioxidant capacity of wild mushrooms compared to commercial mushrooms.12
Mushrooms as Medicine
Views on the consumption of mushrooms varied across the ancient world; while leading Roman medical practitioners were wary of mushroom consumption due to numerous cases of accidental poisoning and excessive consumption of edible mushrooms leading to indigestion by the populace,13 Eastern populations, such as the Chinese, Japanese, and Indians, have long viewed mushrooms as medicinal aids.14,15 Today, the health benefits of various mushrooms are known to include antioxidant, prebiotic, antihypertensive, anti-inflammatory, antiviral/antimicrobial, neuroprotective, hepatoprotective, and antitumor/anticancer effects, among others.16–18
Cardiovascular and metabolic health. Research suggests that edible mushroom consumption may favorably alter metabolic markers (e.g. cholesterol, triglycerides) and reduce blood pressure,19–21 although the effects can differ depending on the mushroom. For example, among spontaneously hypertensive rats (a common animal model of hypertension and cardiovascular disease) in one study, those fed maitake mushrooms (Grifola frondose) experienced a decrease in their total cholesterol level compared to the control group, while those fed shiitake mushrooms (Lentinus edodes) experienced a reduction in their free cholesterol level.20 There was no difference in the plasma triglyceride or phospholipid levels between the experimental groups; however, shiitake consumption resulted in a decrease in both very-low-density lipoprotein (“bad”) cholesterol and high-density lipoprotein (“good”) cholesterol compared to the control group, while maitake consumption elicited a decrease in very-low-density lipoprotein (“bad”) cholesterol only.20 Some of the same investigators reported in another study that the blood pressure of spontaneously hypertensive rats was significantly reduced following eight weeks of maitake mushroom consumption, but this effect was not true with shitake consumption. Moreover, although the investigators affirmed the reduction in plasma-free cholesterol levels and reported reductions in triglyceride and phospholipid levels with shiitake intake, they did not observe reductions in either total or free cholesterol levels or triglyceride and phospholipid levels with the consumption of maitake mushrooms.21 Keeping in mind all these findings and that free cholesterol has cytotoxic effects (which may be mitigated by high-density lipoprotein cholesterol),22 the intake of both mushrooms (ensuring variety) rather than either alone, in combination with other healthy foods (i.e., those that increase high-density lipoprotein cholesterol and phospholipid concentrations on their own), may lead to the best outcome.
Collectively, edible mushrooms appear to support glucose control by a variety of mechanisms, including inhibiting glucose absorption, protecting β-cells (which produce and release insulin in the pancreas) from damage, increasing insulin release, and regulating different relevant pathways in the body.23 In Type 2 diabetic C57BL/6 mice (which carry a genetic predisposition to develop Type 2 diabetes), oral administration of 250 or 500mg/kg of chaga mushroom (Inonotus obliquus) extract significantly alleviated insulin resistance, with a dose–effect relationship noted within a certain range; indeed, the authors reported that the 500mg/kg dose of extract achieved an effect similar to that of the diabetes drug metformin.24 Along these lines, oral administration of 900mg/kg of chaga mushroom in another study led to reductions in fasting blood glucose levels, an improved glucose-tolerance ability, an increased hepatic glycogen level (to better prevent high blood glucose levels), and ameliorated insulin resistance in a Type 2 diabetic mouse model induced by a high-fat diet and streptozotocin (a compound with preferential toxicity toward pancreatic β-cells) compared to diabetic control mice.25
Immune function. Research has attributed the beneficial effects of edible mushrooms on the immune system to their ability to modulate different cytokine responses. In cancer, maitake, Ganoderma lucidum (reishi), Cordyceps sinensis, and Trametes versicolor (turkey tail) may increase the production of T helper (Th)1 cytokines, such as interferon-γ, which activate death receptors on the surfaces of tumor cells to help Th1 cells locate and kill them.26 Edible Agaricus, maitake, reishi, Cordyceps, and turkey tail mushrooms may also downregulate Th2 cytokines, which reduce Th1 cytokine concentrations, thus showing an additional benefit in treating cancer by maintaining higher concentrations of tumor-destroying Th1 cells.26
Edible mushrooms may also increase the therapeutic efficacy of mainstay treatments for cancer.26 During chemotherapy, chemotherapeutic agents penetrate and accumulate in tumor cells to induce cell cycle arrest and apoptosis; as such, some edible mushrooms, such as Agaricus spp., may help drugs such as doxorubicin to accumulate intracellularly at greater doses, increasing their therapeutic efficacy.27 Other edible mushrooms, when combined with such drugs, may help to inhibit tumor growth; one study concluded that administering an extract of Cordyceps sinensis in combination with cisplatin could inhibit tumor growth,28 and another determined that the combination of polysaccharide K (a derivative of turkey tail mushrooms) and trastuzumab reduced cell growth in colorectal tumors by 96 percent.29 Mushrooms may also minimize associated undesirable side effects of chemotherapy and radiation therapy, such as nausea, bone marrow suppression, anemia, and insomnia.30
Other research has examined mushroom intake for managing inflammatory conditions. According to a literature search, the various bioactive molecules found in mushrooms, including peptides, polysaccharides, terpenes, sterols, fatty acids, and phenols, may inhibit major proinflammatory biomarkers and associated pathways, thus exerting anti-inflammatory effects.31 Mushrooms such as chaga,34 maitake,35 and reishi36 also seem to have antiallergic effects by inhibiting the process of mast cell degranulation (during which mast cells release mediators, such as histamine).
Certain edible mushrooms may also combat viral infection by preventing viral entry or replication and stimulating immune cell responses. Polysaccharides from Agaricus blazei Murrill, for example, were found to reduce the cytopathic effects of Western equine encephalitis virus, herpes simplex virus (HSV), and poliovirus in Vero cells (a lineage of cells derived from kidney epithelial cells extracted from an African green monkey).37 A sulfated derivative of a polysaccharide from Agaricus brasiliensis Fr. suppressed HSV-1 and HSV-2 cell attachment, cell penetration, and intracellular spread in vitro.38 Interestingly, the sulfated derivative in question also displayed a synergistic antiviral effect against HSV when combined with the antiviral drug acyclovir, suggesting the potential of combining edible mushrooms with antiviral medications to improve treatment effects.38
Neuroprotection and neuroregeneration. According to investigators, edible mushrooms could play a role in the prevention39 and treatment40,41 of dementia, with various mushroom species displaying the potential to reduce or inhibit the production of beta-amyloid and phosphorylated tau.42 However, mushroom consumption may also help to limit or prevent more general cognitive decline: among 663 participants 60 years of age or older in the Diet and Healthy Aging study in Singapore, those who consumed greater than two portions (>300g) of mushrooms per week had reduced odds of having mild cognitive impairment independent of age, sex, education, cigarette smoking, alcohol consumption, hypertension, diabetes, heart disease, stroke, physical activities, and social activities.43 Similarly, greater mushroom intake was associated with better scores on certain cognitive performance tests among adults 60 years of age or older from the 2011–2014 U.S. National Health and Nutrition Examination Survey.44 Along these lines, a study from western Norway that recruited elderly participants (70–74 years) from the general population confirmed a linear increase in the dose–response association between mushroom consumption and cognitive test performance.45
The neuroprotective effects of edible mushrooms may be attributable to the amino acid ergothioneine, which the human body cannot synthesize itself but can source from certain foods, including mushrooms. Notably, however, whole-blood concentrations of ergothioneine were found to decline significantly after 60 years of age,46 and ergothioneine levels were lower in individuals with mild cognitive impairment (plasma)46 or Parkinson’s disease (serum)47 compared to age-matched healthy individuals. Some edible mushrooms, such as Hericium erinaceus (lion’s mane), contain compounds that may also boost hippocampal memory by encouraging nerve growth.48 The antioxidants in mushrooms may also help to control oxidative stress levels and maintain antioxidant defenses to prevent age-related neurodegeneration.53
Antioxidation. Mushrooms contain both primary and secondary antioxidants as well as compounds with antioxidant properties that act as cell signals and/or inducers, leading to alterations in gene expression that activate enzymes to eliminate reactive oxygen species.50 Certain mushrooms also inhibit lipid peroxidation, a process in which reactive oxygen species trigger the oxidative deterioration of lipids.55 One study determined that mushrooms contain unusually high amounts of ergothioneine and another antioxidant, glutathione, although the levels vary between species: among 13 species tested, maitake (2.41mg/g of dry weight) and Agrocybe aegerita (1.92mg/g of dry weight) mushrooms contained the most glutathione, and Boletus edulis (7.27mg/g of dry weight) and Pleurotus citrinopileatus (3.94mg/g of dry weight) mushrooms contained the most ergothioneine.52
Mushrooms also contain different amounts of other antioxidants, including phenolics, flavonoids, glycosides, polysaccharides, tocopherols, carotenoids, vitamins, minerals, and ascorbic acid.50 In a study from Ethiopia, testing of two cultivated (Pleurotus ostreatus and shitake) and five wild (Laetiporus sulphureus, Agaricus campestris, Termitomyces clypeatus, Termitomyces microcarpus, and Tapura letestui) mushroom species indicated that, among them, Agaricus campestris exhibited significant antioxidant potential due to having the highest levels of multiple phenolic compounds, including ferulic acid, gallic acid, and p-hydroxybenzoic acid.53 In another study investigating hot water extracts of Agaricus, Antrodia, Auricularia, Coprinus, Cordyceps, Hericium, Grifola, Ganoderma, Lentinus, Phellinus, and Trametes mushrooms, researchers reported concentrations of polyphenolic compounds and polysaccharides to be responsible for their high antioxidant potential, with Ganoderma mushrooms exhibiting the greatest antioxidant potential.54 In another study, among 16 of the most popular edible species of wild-growing mushrooms, Boletus chrysenteron and Boletus edulis had high polyphenol contents and antioxidant activity.55
Supplementing with Mushrooms
As an alternative to consuming mushrooms during meals, mushroom supplements are available and often combine multiple mushrooms that are heat-treated and milled to disrupt the chitinous cell wall matrix and increase the surface area for digestion and absorption.56 When choosing a mushroom supplement, however, one may need to consider whether the mushroom mycelium (a web of fibers found underground) or the fruiting body (the cap and stalk) provides better nutrition, as different supplement companies opt to include one, the other, or both.56 Ultimately, to secure one’s preferred nutrient profile, the choice between a mycelium or fruiting body supplement may depend on the mushroom: one study comparing the antioxidant properties of commonly cultivated mushrooms between in-vivo (fruiting body) and in-vitro (mycelium) samples determined that the mushroom species with the greatest antioxidant potential was the brown Agaricus bispous, while, among the mycelium samples, shiitake mushrooms showed the highest antioxidant activity.57 Similarly, other studies reported that the mycelium of Pleurotus ostreatus had greater concentrations of ergosterol and phenolic compounds than the corresponding fruiting body,58 while fruiting bodies of Agaricus bisporus, when compared to both farm (old mycelium) and in-vitro (young) mycelium, contained higher levels of different phenols and ergothioneine.59 In other cases, both parts of the same mushroom may contain unique nutrients: take, for example, lion’s mane, where hericenones were isolated from the fruiting body but erinacines were isolated from the mycelium.60
A Note of Caution
Like other foods, edible mushrooms should be consumed after being properly prepared. Cutaneous reactions (e.g., shitake dermatitis61,62) have been documented following the ingestion of raw or undercooked mushrooms. Raw Agaricus mushrooms also contain agaritine,63,64 a hydrazine-derivative mycotoxin with carcinogenic properties in which concentrations may be reduced—although not removed entirely—by exposing the mushrooms to heat.63 Similarly, Agaricus bisporus and another edible mushroom, Gyromitra esculenta, contain hydrazine analogs, which were found in an animal study following administration in drinking water continuously for life to directly or indirectly (by way of their derivatives) to cause tumors in various tissues in Swiss mice and Syrian (golden) hamsters.65 Of course, serious anaphylactic reactions can occur in susceptible individuals following the consumption of even the most commonly eaten edible mushrooms.66 Finally, mushroom supplementation should be monitored in individuals with more complex health conditions; for example, authors of a case series report of three Japanese patients with cancer suggested a causal relationship between the patients’ severe hepatic damage and their use of Agaricus blazei extract as alternative medicine.67
Editor’s note. Please discuss the consumption of mushrooms or mushroom supplements with your primary care practitioner.
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