Coprinus

Coprinus comatus (Shaggy Ink Cap / Shaggy Mane): A Comprehensive Reference

1. Identity and Taxonomy

Botanical and Scientific Classification

Coprinus comatus (O.F.Müll.) Pers. — also called shaggy mane, chicken drumstick mushroom, or lawyer's wig — is a species of fungus reviewed in Phytotherapy Research (2020) for both its functional properties and potential hazards. C. comatus belongs to the phylum Basidiomycota, family Agaricaceae, and the Coprinus genus. In 1780 it was categorised by Otto Friedrich Müller and first named Agaricus comatus; seventeen years later Christiaan Hendrik Persoon transferred the fungus to the Coprinus genus and gave it its current binomial name.

Its specific epithet derives from coma, meaning "hair," hence comatus, meaning "hairy" or "shaggy." Coprinus comatus is the type species for the genus Coprinus. This genus was formerly considered to be a large one with well over 100 species; however, molecular analysis of DNA sequences showed that the former species belonged in two families, the Agaricaceae and the Psathyrellaceae. C. comatus is the best known of the true Coprinus.

Common Names

• Commonly referred to as either the Shaggy Inkcap or the Lawyer's Wig, Coprinus comatus is a large and conspicuous edible (when young and fresh) fungus.
• In America, the terms Inky Cap or Inky-cap are most commonly used, while Lawyer's Wig and Shaggy Mane are common names used most often in the USA.
• It is also called the "chicken drumstick mushroom" in some Asian markets.

Morphology and Distribution

The young fruit bodies first appear as white cylinders emerging from the ground, then the bell-shaped caps open out. The white caps are covered with scales (the origin of its common names). The gills beneath the cap are white, then pink, then turn black. This mushroom is unusual because it will turn black and dissolve itself in a matter of hours after being picked or depositing spores. The cap ranges from 4–8 cm in width and 6–20 cm in height. It is mostly white with shaggy scales, which are more pale brown at the apex. The free gills change rapidly from white to pink, then deliquesce (melt) into a black liquid filled with spores — hence the "ink cap" name.

Widespread and common in Britain and Ireland, Coprinus comatus is also found throughout mainland Europe, from Scandinavia down to the southern edge of the Iberian Peninsula and the shores of the Mediterranean. It also occurs in North America. In Asian countries, C. comatus is approved as an edible mushroom and often cultivated for consumption, whereas in many other countries, although it is widespread, it is unrecognised and not used.

Common Preparations and Forms

A unique feature of this species is that it is edible only when young, as older fruiting bodies undergo autolysis. In research settings and as a dietary supplement, C. comatus is encountered in several forms. A 96% ethanol extract of the C. comatus fruiting body has been reported to comprise approximately 4.1% fat, 11.4% phenols, 47.8% polysaccharides, 19.9% protein, 0.6% amino acids, 3.9% nucleic acids, and 7.2% fibre. Studies have also used water extracts, ethyl acetate fractions, hexane fractions, chloroform fractions, fermented mycelial preparations, culture-liquid crude extracts, and polysaccharide isolates administered by various routes. Members of the genus Coprinus contain up to 25% protein when young, and they are rich in polysaccharides, triglycerides, and essential amino acids.

2. Traditional and Historical Use

Mushrooms have been used for centuries not only as food but also in traditional medicine as a source of components with pro-health activity. Edible mushrooms have a long history of use in traditional Chinese or Japanese medicine. Coprinus comatus has been used for medicinal purposes in traditional Chinese medicine, where it is believed to have antitumour, antioxidant, and immunomodulatory properties. Coprinus comatus (Agaricaceae) has been used as an antidiabetic food for centuries, particularly in countries such as China; its bioactive compounds, which include polysaccharides, proteins, alkaloids, terpenoids, sterols, and phenolics, have been shown to have a variety of health benefits.

Higher basidiomycetes mushrooms have been used since ancient times in folk medicine to treat a diversity of diseases, including cancer. The black liquid that the mushroom produces as it dissolves itself after maturity is called "ink." In the past, this ink was used as a writing and drawing material, hence the name "ink cap."

In North American indigenous contexts, the pattern of use for this species has varied by community and time period. Secwepemc plant specialist Mary Thomas noted that her people have eaten more kinds of mushrooms recently than they did in the past, noting that shaggy manes (Coprinus comatus) were not eaten traditionally, but within the past few decades she and others liked to pick them and fry them up for a meal.

3. Key Constituents and Active Compounds

Polysaccharides and Beta-Glucans

Polysaccharides extracted from water extracts of C. comatus fruiting bodies have been analysed by NMR spectroscopy and shown to contain disaccharide α,α-trehalose, β-D-glucans, and a lower molecular-mass pentasaccharide-repeating α-L-fuco-α-D-galactan. Studies have found that crude polysaccharides of C. comatus have strong antioxidant effects. GC analysis has confirmed two main polysaccharide components (Ccp-I-A, Ccp-I-B) from the fruiting bodies, composed mainly of mannose, glucose, and galactose. Polysaccharides from C. comatus have hypoglycaemic activity attributed partly to inhibition of non-enzymatic glycosylation (NEG).

Comatin

A compound designated comatin — an inhibitor of the non-enzymatic glycosylation (NEG) reaction — was isolated from Coprinus comatus fermentation broth by macroporous resin separation, followed by chromatographic purification using a C18 reversed-phase column. The compound was identified as 4,5-dihydroxy-2-methoxy-benzaldehyde by high-resolution MS, IR, NMR, and UV analyses. Comatin has been widely reported as a novel hypoglycaemic substance.

Ergothioneine

C. comatus contains ergothioneine, a thiol with antioxidant activity. Ergothioneine exhibits antimutagenic, chemo- and radioprotective activity. In an in vivo study on a hairless mouse model, ergothioneine isolated from Coprinus comatus significantly reduced the number of DNA lesions and inhibited inflammation caused by UV-B radiation. Ergothioneine from C. comatus at 100 μM concentration reduced the activity of myeloperoxidase (MPO) to 0%. Extract from C. comatus at 1 mg/mL showed the same effect. C. comatus also strongly decreased (almost 100%) the activity of the inflammatory marker 8-bromo-2'-deoxyguanosine (8-BrdG) at 1,000 μM of ergothioneine and 1 mg/mL concentration of C. comatus extract.

Vanadium and Trace Elements

Fermented Coprinus comatus is rich in trace elements, including vanadium, chromium, zinc, magnesium, copper, iron, and nickel. Vanadium in particular has attracted scientific attention for glycaemic effects. In fermented mushroom of C. comatus, vanadium at lower doses in combination with C. comatus induced significant decreases of blood glucose and HbA1c levels in hyperglycaemic mice.

Laccases

A novel laccase was isolated and purified from fermentation mycelia of Coprinus comatus through multi-step chromatography. The purified enzyme was a monomeric protein with a molecular weight of 64 kDa, possessing a unique N-terminal amino acid sequence of AIGPVADLKV. This laccase potently suppressed proliferation of tumour cell lines HepG2 and MCF7, and inhibited HIV-1 reverse transcriptase (RT) with IC₅₀ values of 3.46 μM, 4.95 μM, and 5.85 μM, respectively.

Lectins and Proteins

Lectins are proteins or glycoproteins capable of selective binding with membrane carbohydrates of different cell types, fulfilling an important role in the regulation of the immune system. Lectins promote the cell adhesion process, and certain lectins have been found to contribute to the activation of lymphocytes, whereas others possess strong anti-proliferative properties. The broader genus of coprinoid mushrooms has been documented to contain bioactive compounds including sesquiterpenes, proteins, lectins, phenolics, polysaccharides, fatty acids, and enzymes (proteases) possessing medicinal properties including antitumour, antibacterial, antifungal, antioxidant, mitogenic, antiprotozoal, and hypoglycaemic activities.

Other Phenolic Compounds and Alkaloids

The total phenolic content, ergothioneine, and antioxidant activity (AOA) are particularly notable in C. comatus. The bioactive alkaloids in C. comatus function as DPP-4 inhibitors by enhancing the uninterrupted flow of GLP-1 and accelerating phosphatidylinositol (PI) 3-kinase (PI-3K) activity, which enhances insulin biosynthesis and cell proliferation.

4. Scientific Evidence by Area of Activity

4.1 Blood Glucose Regulation and Antidiabetic Effects

The antidiabetic potential of C. comatus is the most extensively studied area of its pharmacological activity. Multiple mechanistic pathways have been identified in preclinical models.

Comatin and Non-Enzymatic Glycosylation Inhibition: The hypoglycaemic effect of comatin on both normal and alloxan-induced diabetic rats was investigated. The blood glucose concentration of normal rats treated with comatin at 80 mg/kg body weight was reduced from 5.14 mM to 4.28 mM in 3 hours. The concentrations of fructosamine, triglycerides, and total cholesterol in induced-diabetic rats were significantly decreased. These results indicated that comatin can maintain a low level of blood glucose and improve glucose tolerance.

Vanadium-Containing Fermented Preparations: The hypoglycaemic activity of fermented mushroom of Coprinus comatus rich in vanadium (CCRV) was studied using alloxan- and adrenalin-induced hyperglycaemic mice. The blood glucose and HbA1c of the mice were analysed. After administration of CCRV, blood glucose and HbA1c of alloxan-induced hyperglycaemic mice decreased (p<0.05, p<0.01), ascension of blood glucose induced by adrenalin was inhibited (p<0.01), and the sugar tolerance of normal mice was improved. In this study, homogenised fermented C. comatus fruiting body was used in a culture medium containing sodium metavanadate (NaVO₃) (CCRV). The level of hepatic glycogen was increased by the use of this combination. In mice fed CCRV, glycogen level was 27.6 ± 5.2 mg/g — higher than in diabetic mice (14.1 ± 3.8 mg/g) and comparable to normal mice (24.1 ± 4.3 mg/g). Damaged pancreatic β-cells were perceptible in diabetic mice, but CCRV-fed mice did not show loss of pancreatic cells.

Trace-Element Comparison Study: The effect of fermented Coprinus comatus rich in trace elements — including vanadium, chromium, zinc, magnesium, copper, iron, and nickel — on glycaemic metabolism was studied in alloxan-induced hyperglycaemic mice. Blood glucose, glycohaemoglobin, and glycogen synthesis were analysed. After administration of C. comatus rich in vanadium (CCRV), blood glucose and glycohaemoglobin decreased (p<0.05, p<0.01), glycogen synthesis was elevated (p<0.01), gluconeogenesis in normal mice was inhibited (p<0.01), and sugar tolerance was improved. However, the same result did not occur in other groups. Vanadium at lower doses in combination with C. comatus induced significant effects on glycaemic metabolism in mice.

GLP-1 and DPP-4 Pathway: One study aimed to identify the effect of bioactive compounds of C. comatus extract as an antidiabetic agent linked to glucagon-like peptide 1 (GLP-1) and antioxidant properties in increasing glutathione (GSH) levels. The study used six groups of Wistar rats (n = 24), with groups 2–6 receiving 45 mg streptozotocin/kg body weight; groups 4–6 were also given 250, 500, and 750 mg of C. comatus ethyl acetate extract/kg body weight for 14 days. The bioactive alkaloids in C. comatus have been described as DPP-4 inhibitors that enhance the flow of GLP-1 and accelerate PI-3K activity, enhancing insulin biosynthesis. Previous research confirmed that ethyl acetate extract of C. comatus at 500 mg was effective in increasing levels of GLP-1.

Methanolic Extract Study: One study investigated C. comatus methanolic extracts for antidiabetic and antioxidant activity. Sixty male rats were divided into five groups. On the basis of results obtained, the 400 mg/kg body weight C. comatus extract significantly reduced fasting blood glucose levels by up to 22.86% in the treated group.

Evidence Strength: All published antidiabetic evidence for C. comatus to date derives from animal models (rodents) and in vitro experiments. No randomised controlled human clinical trials on C. comatus specifically for glycaemic control have been identified in the peer-reviewed literature. The preclinical findings are consistent across multiple research groups and mechanistic pathways, but translation to human benefit is unproven.

4.2 Antioxidant Activity

Ethanolic and hot-water extracts of fruiting bodies and mycelium, as well as fermented filtrate of C. comatus, have shown high antioxidant activity (AOA). Polysaccharides from C. comatus affect hepatic and mitochondrial antioxidant enzymes. Treatment with polysaccharides increased activity of hepatic glutathione peroxidase (GSH-Px) by approximately 166.78%, SOD by approximately 83.72%, and catalase (CAT) by approximately 63.12%. Activity of mitochondrial enzymes GSH-Px, SOD, and CAT was increased by C. comatus by approximately 92.00%, 67.03%, and 51.61%, respectively. Song and Du demonstrated medium antioxidant ability of C. comatus polysaccharides compared with other tested mushrooms on assays for superoxide anion radical scavenging activity (SRSA), reducing power (RP), and chelating ability (CA), and weak antioxidant ability on assays for hydroxyl radical scavenging activity (HRSA) and DPPH scavenging ability (DSA).

Evidence Strength: Antioxidant activity has been demonstrated in multiple in vitro and animal studies. As with glycaemic data, no controlled human trials have been published. Evidence is preliminary and largely biochemical.

4.3 Anticancer and Antiproliferative Activity

The IC₅₀ value of the culture-liquid crude extract of C. comatus on MCF7 (breast cancer) cell viability was as low as 76 μg/mL, and the IC₅₀ value of the ethyl acetate extract was 32 μg/mL. Both extracts significantly affected IκBα phosphorylation in a dose-dependent manner. The effect of the ethyl acetate extract was comparable to curcumin, a known NF-κB pathway inhibitor. The ethyl acetate extract inhibited the activity of the IKK complex at close to 90% compared to untreated controls. These results suggest that C. comatus contains potent compounds capable of inhibiting NF-κB function and may act as an antitumour agent.

Treatment of ES-2 ovarian cancer cells with ethyl acetate extract of C. comatus (100 μg/mL medium) for 48 or 72 hours resulted in an increased number of cells in the sub-G1 phase of the cell cycle, an increased number of apoptotic cells, and reduced levels of procaspases -3, -8, and -9. Ethyl acetate extract of C. comatus was concluded to induce apoptosis in ovarian cancer cells (ES-2) via both extrinsic and intrinsic pathways.

Antiandrogenic Activity / Prostate: One study evaluated the antiandrogenic activity of different Coprinus comatus strains in their ability to interfere with androgen receptor (AR) function. The most active extract was C. comatus strain 734 extracted with hexane (CC734-H). This extract was able to inhibit AR-mediated reporter activity, inhibit the proliferation and viability of the LNCaP prostate cancer cell line, and inhibit colony formation of the LNCaP cell line, compared to DU-145, PC-3, and MDA-Kb2 cells. In addition, CC734-H was able to reduce AR levels and prostate-specific antigen gene expression in the LNCaP-treated cell line. The study illustrated the potential of C. comatus as a natural antiandrogenic modulator that could serve in treatment of prostatic diseases.

Anti-Glioma: Research has also assessed extracts of C. comatus alongside other mushrooms as potential therapeutic agents for treatment of cancer including glioma, as referenced in the broader medicinal mushroom literature.

Evidence Strength: All anticancer findings are from in vitro cell-line studies and animal tumour models. More investigations are needed to determine whether the apoptotic effect on cancer cells is accomplished by one active compound or combined activities of different compounds. No human clinical trials evaluating C. comatus for cancer treatment have been published. Evidence is early-stage and mechanistic; clinical relevance is unestablished.

4.4 Hepatoprotective Effects

The aim of one study was to characterise alkalic-extractable polysaccharides (ALPS) from Coprinus comatus and to explore their in vivo antioxidant activities and protective effects on alcohol-induced liver injury. ALPS showed strong antioxidant and anti-inflammatory abilities, markedly low serum enzyme activities, hepatic and serum lipid levels, as well as low hepatic lipid peroxidation levels; ALPS also improved the alcohol metabolism system. These results were confirmed by histopathological section analysis.

One study aimed to investigate the hepatoprotective effects of Coprinus comatus protein (CCP) in a mouse model of acute alcoholic liver injury by examining regulation of gut microbiota dysbiosis. Recent studies have also suggested a relationship between C. comatus and gut microbiota, in which the hypoglycaemic mechanism of C. comatus is involved, at least in part.

Evidence Strength: Hepatoprotective data are exclusively from mouse models. No human clinical data exist. Evidence is preliminary.

4.5 Anti-inflammatory Activity

Ergothioneine from C. comatus at 100 μM concentration reduced the activity of MPO (myeloperoxidase, an inflammatory marker) to 0%, and C. comatus extract showed the same effect at 1 mg/mL. C. comatus also strongly decreased (almost 100%) the activity of inflammatory marker 8-BrdG at 1,000 μM ergothioneine and 1 mg/mL extract concentration. These results indicated anti-inflammatory properties of C. comatus.

Evidence Strength: Anti-inflammatory evidence is in vitro only. No human studies on inflammation endpoints have been conducted with C. comatus.

4.6 Antiviral Activity

Purified laccase from C. comatus may cause inhibition of the HIV-1 reverse transcriptase (HIV-1 RT) protein at IC₅₀ = 5.85 μM. HIV-1 RT is an important element in the retroviral life cycle because it participates in the synthesis of double-stranded DNA from a single RNA genome. Consequently, HIV-1 RT has been suggested as a main antiviral drug target, and in the future the laccase could potentially be used in support of AIDS treatment.

Evidence Strength: Antiviral data are exclusively from in vitro enzymatic inhibition studies. No animal or human studies have been reported. This finding is highly preliminary.

4.7 Antimicrobial Activity

The antimicrobial properties of C. comatus are among its well-documented bioactive properties in the published literature. The broader coprinoid family has been reported to contain bioactive compounds possessing antibacterial and antifungal activities.

Evidence Strength: Antimicrobial evidence rests primarily on in vitro studies. Clinical relevance is not established.

4.8 Nematocidal Activity

Coprinus comatus is a nematophagous fungus capable of killing and digesting the nematode species Panagrellus redivivus. It does so by making a structure called a "spiny ball," a burr-like structure assembled with a large number of tiny tubes. Nematodes added to C. comatus cultures grown on nutrient agar become inactive within hours. Electron microscopy shows that C. comatus infects P. redivivus by producing penetration pegs from which hyphae colonise nematode bodies. Within days, the infected nematode is digested and consumed by mycelial hyphae. This may be a mechanism to help the fungus thrive in nitrogen-poor environments.

5. Body Systems and Health Areas Associated with Coprinus comatus

Various studies show physiological activities of C. comatus across multiple domains, including antioxidant, anticancer, antiandrogenic, hepatoprotective, acetylcholinesterase inhibitory, anti-inflammatory, antidiabetic, antiobesity, antibacterial, antifungal, antinematode, and antiviral activities.

• Endocrine / Metabolic System: Blood glucose regulation and antidiabetic effects via comatin-mediated NEG inhibition, vanadium-associated insulin-mimetic effects, DPP-4 inhibition, and enhancement of GLP-1 signalling.
• Immune System: Immunomodulatory and antitumour effects; lectin-mediated lymphocyte activation; polysaccharide-driven immune signalling.
• Hepatic System: Protection against alcohol-induced liver injury via polysaccharide-mediated antioxidant enzyme upregulation and lipid-lowering effects.
• Oncological / Antiandrogenic: Androgen receptor modulation in prostate cancer cell lines; NF-κB pathway inhibition in breast cancer; caspase activation in ovarian cancer cells (all in vitro).
• Skin / Inflammatory: Ergothioneine-mediated reduction of UV-B-induced DNA lesions and myeloperoxidase activity.
• Neurological (Preliminary): Research has assessed Coprinus comatus inhibitory capacity against acetylcholinesterase (AChE) and butyrylcholinesterase (BChE), enzymes linked to Alzheimer's disease.

6. Dosage Forms and Reported Dosages

No standardised human dosage for Coprinus comatus supplements has been established in clinical trials. The following dosages were used in the preclinical (animal) studies reviewed in the literature:

• Comatin administered at 80 mg/kg body weight in rats reduced blood glucose from 5.14 mM to 4.28 mM within 3 hours.
• Streptozotocin-induced diabetic rats received 250, 500, and 750 mg of C. comatus ethyl acetate extract per kg body weight for 14 days.
• In an alloxan-induced diabetic rat study, animals received 200 and 400 mg/kg body weight mushroom extract; the 400 mg/kg dose reduced fasting blood glucose by up to 22.86%.
• In breast cancer cell studies, IC₅₀ values for culture liquid crude extract on MCF7 cell viability was 76 μg/mL, and for the ethyl acetate extract was 32 μg/mL.
• In ovarian cancer cell experiments, ES-2 cells were treated with 100 μg/mL ethyl acetate extract for 48 or 72 hours.
• In vitro, the laccase from C. comatus mycelia inhibited HIV-1 reverse transcriptase with an IC₅₀ of 5.85 μM.

These figures are from preclinical research and are not established human doses. No pharmacopoeial monograph specifying a human dose for C. comatus has been identified in the literature reviewed.

7. Safety Considerations and Notable Interactions

Coprine and Alcohol: Critical Species Clarification

Coprinopsis atramentaria (formerly classified within Coprinus) is edible but contains coprine, which causes a negative disulfiram-like reaction related to consuming this Basidiomycota with ethanol. Coprine is chemically N5-(1-hydroxycyclopropyl)glutamine. It causes inhibition of aldehyde dehydrogenase in the liver, which increases the level of acetaldehyde in the blood after alcohol consumption.

Coprine has been isolated from Coprinopsis atramentaria (formerly Coprinus atramentarius); however, other Coprinus species such as the common edible Coprinus comatus (Shaggy Mane, Lawyer's Wig) do not contain coprine. Other Coprinus species do not contain coprine, including the edible Coprinus comatus.

Despite this, a practical hazard remains because of potential misidentification. Coprinus comatus can sometimes be confused with Coprinopsis atramentaria, which is found in similar places and has a cap similar to that of C. comatus. When young and fresh, Coprinus comatus is generally considered edible and does not cause coprine reactions, but confusion with the Common Ink Cap is possible. If alcohol is consumed up to approximately 72 hours after the ingestion of species containing coprine, alcohol intolerance occurs, similar to antabus effects, with an intense indisposition for several hours, including decreased blood pressure, elevated heart activity, flushing, nausea, vomiting, and headache.

Some sources have raised a note of caution even for C. comatus itself in combination with alcohol: Consuming Coprinus comatus in combination with alcohol can cause a reaction that leads to unpleasant symptoms, such as nausea and flushing, so caution is still advised. However, the established scientific position is that C. comatus itself does not contain coprine.

Heavy Metal Bioaccumulation

C. comatus young fruiting bodies are edible. Eating them when foraged from urban places can provide to a consumer mercury at a relatively high dose; the absorption rate of mercury compounds contained in ingested mushroom is an unresolved question. Macrofungi are well known for their ability to efficiently absorb various metallic elements and metalloids from substrates and to sequester them in their fruiting bodies. This bioaccumulation potential is highly dependent on the quality of the growing substrate and soil, making the geographic provenance and cultivation conditions of commercial preparations a relevant safety consideration.

Allergenic Potential in Atopic Individuals

Coprinus comatus may cause skin reactions in patients with atopic dermatitis and atopic predisposition. One study showed that in 32% of patients with atopic dermatitis, C. comatus induced delayed-type reactions after an atopic patch test (containing 5 mg protein from cap per 1 g vaseline or 1.35 mg spore protein per 1 g vaseline). After 48 and 72 hours, negative skin test reactions were observed in the non-atopic control group.

Absence of Human Interaction Data

There are no published human clinical trials identifying drug interactions for C. comatus preparations. The theoretical concern for additive hypoglycaemic effects when combined with antidiabetic medications arises from preclinical data demonstrating blood-glucose-lowering activity, but this has not been formally evaluated in human studies. Similarly, given the laccase's in vitro inhibition of HIV-1 reverse transcriptase, no human pharmacokinetic or interaction data exist.

8. Summary of Evidence

Various studies show many of the physiological activities of C. comatus, including antioxidant, anticancer, antiandrogenic, hepatoprotective, acetylcholinesterase inhibitory, anti-inflammatory, antidiabetic, antiobesity, antibacterial, antifungal, and antinematode activities. However, the great majority of this evidence derives from in vitro cell culture experiments and rodent models. Coprinus comatus has not been evaluated in a randomised, placebo-controlled human clinical trial for any indication as of the sources reviewed. Existing preclinical data are mechanistically diverse and internally consistent across research groups, which supports continued investigation, but clinical efficacy and human safety profiles remain to be established.

References

• Nowakowski P, et al. "The two faces of Coprinus comatus—Functional properties and potential hazards." Phytotherapy Research, 2020. PMC7754439
• Dotan N, Wasser SP, Mahajna J. "The Culinary-Medicinal Mushroom Coprinus comatus as a Natural Antiandrogenic Modulator." Integr Cancer Ther, 2011. PMID 21147815
• Rouhana-Toubi A, et al. "Coprinus comatus substances interfere with H2O2 induction of the NF-kappaB pathway in MCF7 breast cancer cells." Int J Med Mushrooms, 2011. PMID 22135900
• Rouhana-Toubi A, Wasser SP, Fares F. "The Shaggy Ink Cap Medicinal Mushroom, Coprinus comatus Extract Induces Apoptosis in Ovarian Cancer Cells." Int J Med Mushrooms, 2015. PMID 26854099
• Li W, et al. "The Saggy Ink Cap Medicinal Mushroom, Coprinus comatus Protein Attenuates Acute Alcoholic Liver Injury in Association with Changes in the Gut Microbiota of Mice." Int J Med Mushrooms, 2021. PMID 34347998
• Han C, et al. "Hypoglycemic activity of fermented mushroom of Coprinus comatus rich in vanadium." Journal of Trace Elements in Medicine and Biology, 2006.
• Zhou S, et al. "Comparison of hypoglycemic activity of trace elements absorbed in fermented mushroom of Coprinus comatus." PubMed, 2009. PMID 19283342
• Ratnaningtyas NI, et al. "Ethanol extract of the mushroom Coprinus comatus exhibits antidiabetic and antioxidant activities in streptozotocin-induced diabetic rats." Pharmaceutical Biology, 2022. PMC9186368
• Medicinal Mushrooms for Treatment of Type 2 Diabetes: An Update on Clinical Trials. Int J Med Mushrooms, 2020.
• Zhao H, et al. "The antioxidant activities of alkalic-extractable polysaccharides from Coprinus comatus on alcohol-induced liver injury in mice." Scientific Reports, 2018. PMC6076309
• Zhao S, et al. "A Novel Laccase with Potent Antiproliferative and HIV-1 Reverse Transcriptase Inhibitory Activities from Mycelia of Mushroom Coprinus comatus." Biomed Res Int, 2014. PMC4164474
• Mercury bio-extraction by fungus Coprinus comatus: a possible bioindicator and mycoremediator of polluted soils? PMC4846694
• Comparative overview of antioxidative properties and phenolic profiles of Coprinus comatus and Coprinellus truncorum. PMC5306037
• Coprinus comatus — an overview. ScienceDirect Topics.
• Coprinus (toxicology overview). ScienceDirect Topics.
• Coprinus comatus. Wikipedia (taxonomy and morphology section).
• First Nature: Coprinus comatus, Shaggy Inkcap or Lawyer's Wig mushroom.
• 'Frog's umbrella' and 'ghost's face powder': the cultural roles of mushrooms and other fungi for Canadian Indigenous Peoples. Canadian Journal of Botany, 2022.
• Ratnaningtyas NI et al. "Antidiabetic Effects and Antioxidant Properties of the Saggy Ink Cap Medicinal Mushroom, Coprinus comatus, in Streptozotocin-Induced Hyperglycemic Rats." Int J Med Mushrooms, 2021.
• Antidiabetic and Antioxidant Potential of Coprinus comatus in Alloxan-Induced Diabetic Rats Models. Journal of Population Therapeutics and Clinical Pharmacology, 2023.
• Coprine – Mushroom Toxin with Alcohol Reaction. RuggedBears, 2025.