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Substrate Matters

Updated: 8 hours ago

HR (Host Relations) for mushrooms

Ganoderma oregonense 8/2025
Ganoderma oregonense 8/2025

Medicinal mushrooms contain numerous bioactive compounds that influence a variety of physiological processes. Some of these compounds are produced de novo, influenced by growing conditions, while others are derived from the medium, substrate, or host that the mycelium colonizes. This paper explores the presence of different constituents in mushrooms based on their growing conditions: innately biosynthesized versus tree-derived compounds in wild mushrooms and biotransformed plant compounds from medicinal plant-enriched substrates. It also examines how various cultivation conditions influence the biosynthesis of desirable endogenous chemicals such as ergothioneine, 𝛽-glucans, cordycepin, terpenoids, hericenones, erinacines and psilocybin.


In The Wild

There is significantly more variability of bioactive compounds in wild mushrooms: due to the stress of the elements and exposure to pathogens, wild mushrooms produce more secondary metabolites than their cultivated counterparts. Secondary metabolites include terpenoid compounds, flavonoids and phenols and are synthesized by mushrooms in response to environmental stressors and pathogens, often independent of the host. Mushrooms are also bioaccumulators and take up minerals and heavy metals from the environment and host they are colonizing (Siwulski 2025). While most mushroom chemistry appears to be innately biosynthesized, in some cases, the host tree may also determine the chemical makeup of the mushroom. 

Primary Metabolites

(Essential for mushroom survival and growth)

Examples

Polysaccharides 

𝛽-glucans, mannans, chitin

Proteins and peptides

Lectins, enzymes

Lipids and fatty acids

Ergosterol

Amino acids

Glutamate, aspartate, tryptophan, ergothioneine

Nucleotides and derivatives

Cordycepin, adenosine


Secondary Metabolites 

(Non-essential for growth, pathogen and environmental defense and adaptation)

Examples

Terpenes and steroids

Triterpenes, sesquiterpenes, diterpenes

Ganoderic acids, hericenones, erinacines

Phenolic compounds and polyphenols

Flavonoids, hispidin, hispolon

Alkaloids and indoles

Psilocybin, psilocin, tryptamine


De Novo Secondary Metabolites

A study of 10 mushroom species in northeast Thailand found that wild mushrooms had significantly higher total antioxidant capacity, total phenolic content and total flavonoid content than cultivated counterparts, suggesting that environmental stressors enhance production of bioactive metabolites (Srikram 2016). Another review confirmed that wild-harvested mushrooms often contain greater levels of phenolics, terpenoids, and flavonoids compared to cultivated mushrooms (Gebre 2023). These studies focused solely on innately biosynthesized mushroom compounds that were not necessarily dependent on resins and other components from the host tree.


Tree-derived Compounds

To date, there has been little investigation regarding fungal accumulation of particular tree-derived phytochemicals in fruiting bodies, as most profiling focuses on endogenous fungal secondary metabolites and not tree-specific substances. There are two primary ways to determine if compounds found in mushrooms originate from the host tree: isotope-labeling tracers and targeted metabolomic studies comparing mushrooms from different host trees to identify tree-derived markers in fruiting bodies. Isotope-labeling tracer studies are lacking, and there are few targeted metabolomic studies. The best (and worst) example of tree-derived compounds bioaccumulating in a mushroom is in the case of Inonotus obliquus (chaga).


Chaga and Birch 


Chaga as we know it is not in fact a mushroom, but a sclerotium, a mass of infected tree tissue colonized by the fungus Inonotus obliquus. This sterile canker contains many innate secondary metabolites as well as birch-derived terpenes (betulin and its oxidized form, betulinic acid). Chaga is a complex example of a fungus taking up compounds from trees because the sclerotium - the part that is traditionally used therapeutically - and the tree matter are largely indistinguishable (Windsor 2025).


This brings up the question: can chaga truly be cultivated? My opinion is no - chaga as we know it can not be cultivated. The cultivated Inonotus obliquus has medicinal value, but it is not the same as chaga, the mixture of decaying birch and mycelial biomass. This relationship seems more similar to a lichen (fungi and algae) although that relationship is mutualistic. Wild chaga can only exist in its sclerotium form because of its parasitic relationship with the birch tree, and therefore the name “chaga” should only refer to this dual-form structure.


Birch bark is rich in lupane-type triterpenes - betulin mostly, and betulinic acid in smaller amounts. Chaga lacks the complete enzymatic pathway to synthesize these triterpenes. The most likely explanation of their presence in chaga is passive absorption or biotransformation of bark-derived betulins into the fungal tissue during colonization of the host wood. Other wood-inhabiting fungi, the famous Fomitopsis betulina, the favorite of Ötzi the Iceman, has also shown trace betulinic acid when harvested from birch trees, suggesting possibly a generalizable phenomenon across polypores (Sułkowska-Ziaja 2018).


A comparison study exploring chaga from different alder and birch species and from different locales  (Estonia and Finland) demonstrated the presence of different bioactive substances depending on the host tree and place of origin. Chaga growing on birch from Estonia contained the most betulin, and surprisingly chaga from Estonian alder contained the most betulinic acid. Ultimately, there were more bioactive compounds - inotodiol, lanosterol, polyphenols and glucans - found from birch-derived chaga, especially birch from Finland, when compared with chaga from alder (Drenkhan 2022). The varying presence of betulin and betulinic acid points to host uptake rather than innate biosynthesis, while the omni-presence of other compounds points to endogenous production, likely varying slightly due to environmental conditions. Betulin and betulinic acid are proposed to be the main anti-mycobacterial components produced by alder and birch and the presence in chaga is likely a defense mechanism of the tree (Li 2015).


A white paper by mycologist Steve Farrar and herbalist Robert Dale Rogers argues that organically cultivated Inonotus obliquus mycelium, often labeled “cultivated chaga”, offers comparable or even superior functional benefits to wild-harvested chaga.The cultivated Inonotus obliquus has the benefit of being more than 15% fungal biomass (often up to 90%), is rich in polysaccharides and antioxidants, lower in oxalates and less likely to contain heavy metal contamination and environmental pollutants (Rogers 2025). This all seems ideal, and illuminates that cultivated Inonotus obliquus has medicinal properties, but completely redefines chaga as fungal mycelium only and excludes the birch tree component, which as stated earlier, is an integral part of the chaga organism. 


If the birch aspect of chaga is so desirable, then perhaps the grower could simply add birch to the substrate and expect higher levels of betulin and betulinic acid. However, researchers have found the opposite effect to be true - when birch-derived Inonotus obliquus mycelium was grown on substrate containing 5% birch, it contained less betulin and betulinic acid than when grown on substrate without birch. Not surprisingly, this study found significantly less betulin and betulinic acid in alder and hornbeam-derived mycelium (Sułkowska-Ziaja 2023).


Overall, information is significantly lacking regarding bioactive compound uptake from host tree to mushroom and the best example we have of this is with chaga, though this is a complicated example. Other polypores well known for their rich terpenoid and phenolic profiles, Fomitospsis spp., Trametes spp. and Ganoderma spp. may take up specific host-tree metabolites into their fruiting bodies or mycelium, but there has yet to be any peer-reviewed research. Further research with isotope-tracing studies is needed to map uptake versus fungal biotransformation.


Therapeutic Applications of Betulin and Betulinic Acid

Betulin and betulinic acid exhibit anti-inflammatory, antiviral, antibacterial and anticancer properties. Betulinic acid has specifically been studied for its selective cytotoxicity toward tumor cells. They also have antioxidant effects and have been shown to support skin health, immune function and overall metabolic balance (Zuco 2002, Fuldo 2008, Lou 2021). 


Comparison of Wild Chaga vs Cultivated Inonotus obliquus

Compound/Class

Wild 

Cultivated

Therapeutic implications

Birch-derived terpenes (betulin and betulinic acid)

Present (Windsor 2025, Rogers 2025, Sułkowska-Ziaja 2023)

Absent (Windsor 2025, 

Rogers 2025)


Present if originally isolated from birch (Sułkowska-Ziaja 2023)

Anti-inflammatory, antiviral, antibacterial and anticancer properties

Phenolic compounds

Present and high diversity (Windsor 2025)

Present, but generally lower diversity (Windsor 2025)

Anti-oxidant, anti-inflammatory

Melanin (pigments)

Present, especially in dark sclerotia (Windsor 2025)

Mostly allomelanin (birch-derived) (Rogers 2025)

Present

Mostly eumelanin (Rogers 2025)

Present, but in lower concentrations (Windsor 2025)


DNA protection against oxidative damage, radioprotective properties

Ergosterol

Present, but typically lower (Windsor 2025, Rogers 2025)


Present and sometimes higher (82% more) (Windsor 2025, Rogers 2025)


Precursor of vitamin D2

Triterpenes (lanostane type - Inotodiol)

Abundant (Windsor 2025)

None detected (Windsor 2025)

Immune-modulating, antioxidant

Polysaccharides 

𝛽-glucans present but low compared with other mushroom species (8-15%)(Windsor 2025)

Endopolysaccharides present (Rogers 2025) High amounts of ɑ-glucans (Windsor 2025

(𝛽-glucans)

Immuno-modulating, prebiotic

Contaminants/heavy metals

Risk due to long-term growth on birch in polluted environments (Rogers 2025)

Very low (Rogers 2025)

Heavy metal toxicity if used long-term

Oxalates

High oxalate content

31mg/g (Rogers 2025)

Low oxalate content 4mg/g (Rogers 2025)

Potential kidney stone risk and other oxalate sensitivity issues


Cultivation

Mushrooms and Medicinal Plant Duos

While research on tree-derived compounds in mushrooms is limited, there is much more evidence for substrate-derived compounds in cultivated mushrooms. In many cases, bioactive compounds from medicinal plants have been shown to be absorbed or biotransformed in the mushroom. This opens up a new frontier for medicinal mushrooms, suggesting a future where different mushroom strains could contain various compounds from medicinal plants and possibly make them more bioavailable.


Shiitake and Astragalus

Astragalus, a potent immune modulator and considered a “qi tonic” in Traditional Chinese Medicine (TCM) can be an influential substrate component (Zheng 2020). Shiitake mushrooms grown on Astragalus membranaceus root powder comprising 5% and 20% of total substrate had total polysaccharides increase by 35-60%. The monosaccharide profile shifted toward astragalus-type rhamnose/galacturonic acid and polysaccharide-rich extracts showed stronger anti-proliferative activity on colon cancer cells compared with shiitake grown on Astragalus-free substrate (tamang 2021). 


Substrate Astragalus concentrations of 5% seemed to take up a similar amount of compounds as the 20% bed, with some minor differences, while these compounds were absent from shiitake grown on Astragalus-free substrate. This analysis confirms the hypothesis that cultivating shiitake on an optimized Astragalus bed facilitates the transfer of compounds from Astragalus to shiitake (Tamang 2021, Balakrishnan 2021).


Therapeutic Applications of Astragalus Polysaccharides

Astragalus polysaccharides are immunomodulators that enhance white blood cell activity - lymphocytes and dendritic cells - and tumor suppression (Shi 2024, Zheng 2020). Preclinical studies have demonstrated anti-tumor, antioxidant and neuroprotective properties. Astragalus polysaccharides may also be useful in neurodegenerative disease via regulation of specific cell signaling pathways - PI3K/AKT/mTOR, JAK/STAT, and Nrf2 (Shi 2024). Clinically, in a phase 2 trial, Astragalus polysaccharides reduced chemotherapy-induced fatigue and improved quality of life in breast cancer patients (Shen 2024). Meta-analyses also support its safe use alongside cancer therapies to enhance immune function (Li 2025).


Oyster mushroom and Chinese formula

Oyster mushrooms cultivated on corncob substrates containing 30–60% residues from Traditional Chinese Medicine (TCM) formulas, Compound Kushen Injection (CKI), Qizhi Tongluo Capsule, and Shenbai Shuxin Capsule, showed significantly higher levels of bioactive compounds - phenolics, flavonoids, terpenoids, and vitamin C - compared to those grown without TCM residues (Jin 2018, 2020). Notably, these mushrooms did not absorb any toxic components from the formulas.


Another study confirmed that using medicinal herb residue-integrated substrate increased polysaccharide and saponin levels in oyster mushrooms grown on substrate containing 80% residue and 20% sawdust. Unfortunately, this study did not identify the exact chemical compounds from the herbs that ended up in the mushrooms, but did show increased levels of polysaccharides and saponins, implying uptake or biosynthesis (Huynh 2025).


Oyster mushroom composition is significantly influenced by the medium on which they are growing, and they are exceptional decomposers capable of breaking down hydrocarbons, pesticides, herbicides, heavy metals, and even plastics (Utilization 2023, Siracusa 2017). These findings suggest that they offer a promising avenue for research into the uptake of bioactive compounds from medicinal plants as well.


Enoki and Green Tea

Enoki mushrooms grown on compost containing 10% green tea powder had significantly more caffeine than enoki without green tea compost. Caffeine levels increased from 3mg/kg dry weight  to 45mg/kg dry weight. However, catechins, the polyphenols abundant in green tea, did not transfer, likely due to their large molecular size (Lee 2008).


Oyster Mushroom and Spent Coffee Grounds

Mushrooms not only absorb compounds from their environment, but they also frequently transform them into new metabolites and other compounds. For example, Pleurotus mycelium, grown on spent coffee ground-integrated substrate, demethylates caffeine into theobromine, and similar to enoki, caffeine has been detected in the mushroom cap after harvest. While caffeine and its metabolites have been found in fruiting bodies, it's unclear if these metabolites are simply transferred from the mycelium or if they are altered within the fruiting body itself. Eating oyster mushrooms grown on spent coffee grounds are unlikely to give you a boost though - to get the caffeine equivalent of one espresso shot from oyster mushrooms grown on spent coffee grounds, you'd need to consume about 250 kg (550lbs) of fresh mushrooms (Carrasco 2019).


Therapeutic Applications of Caffeine

Caffeine has significant cognitive benefits, it enhances physical performance and has been shown to improve mood, especially in the morning. Caffeine improves reaction time, attention and mood by blocking adenosine receptors and increasing dopamine and norepinephrine levels  (Schepicii 2020, Čižmárová 2025). Observational studies have shown that caffeine may also lower dementia risk (Rodak 2021). Human trials also confirm that caffeine can improve endurance and power output, improving overall athletic performance (Guest 2021). In a study of over 200 young adults, caffeine consumed within the first few hours of waking up significantly (and unsurprisingly) boosted positive emotions (Hachenberger 2025).


Ganoderma and Ginseng

When the “Mushroom of Immortality” derives nutrients from a substrate mixture containing the “King of Herbs” the result is possibly the elixir of life. Probably not a true panacea, but likely remarkable medicine. Reishi grown on substrate containing Panax ginseng root powder contained minor and rarer forms of ginseng-specific compounds (ginsenosides). Furthermore, the overall concentration of ginsenosides in fungal biomass increased as did synergistic antioxidants and anti-inflammatory activity (Bai 2025).


Therapeutic Applications of Ginsenosides 

There are over 130 human clinical trials using ginseng that have been registered, many of which are randomized and placebo-controlled (He 2018). Ginsenosides, saponins derived from ginseng, have a multitude of potential benefits - neuroprotective, cardiometabolic, and anti-inflammatory effects to name a few (Lee 2014). They have been shown to enhance learning and memory in animals, and preliminary clinical trials have also shown improvements in cardiovascular and metabolic health outcomes (Feng 2022, Li 2023).


Mushrooms and Soybean meal

Soy-based substrates are commonly used to grow oyster, shiitake and other medicinal culinary mushrooms. Unfortunately, there is not yet evidence of fruiting bodies taking up phytoestrogenic isoflavones from soybean meal substrate, but Pleurotus ostreatus, Hericium erinaceus, and Flammulina velutipes mycelium demonstrated enzymatic hydrolysis of isoflavone glycosides into their more bioavailable and desirable aglycone form - genistein and daidzein (Wang et al. 2023).  In plants, isoflavones most often exist as glycosides (they are bound to a sugar molecule). When the sugar is removed, via hydrolysis by mycelium, or in the intestines of humans (largely dependent on microbiome composition), an aglycone is formed, a smaller and significantly more bioavailable molecule. This research suggests that not only can mycelium take up molecules from plants, but also has potential to make them more bioavailable for human health. The most well known example of this is tempeh, when soy is colonized and fermented by Rhizopus oligosporus fungus tempeh consequently contains more bioavailable isoflavones in their aglycone form.


Therapeutic Applications of Isoflavones

Soy is a controversial plant for many reasons - it is a major monocropped plant whose cultivation plays a major role in rainforest destruction and contains a significant amount of phytoestrogenic isoflavones. Soy monocropping is in fact terrible for the environment, but phytoestrogenic isoflavones are not to be feared, and are therapeutic in conditions where estrogen is dominant or deficient. Soy isoflavones bind to estrogen receptors, particularly ER-β, where they exert modest estrogenic or anti-estrogenic effects depending on endogenous hormone levels. This selective receptor activity helps balance estrogen dominance or deficiency, supporting hormonal homeostasis (Canivenc 2023).


Patent Examples

As illustrated, substrate composition makes a difference in bioactive compounds present in mushrooms. In addition to different substrates impacting the concentration of various desirable compounds, mushroom growth medium and environment also impacts mushroom yield and rate of growth. Considering all of this, finding the perfect substrate and environment is highly desirable and so naturally, companies patent these mixtures to protect their cultivation formula. A few examples of patented substrate mixtures that change the chemical composition of mushrooms are ginseng and cordyceps and purple corn and reishi.


Cordyceps and Ginseng

Patent KR100610983B1 – Cultivation of Cordyceps sinensis using ginseng medium (filed 2004) uses a substrate blend replacing silkworm pupae with 25-100% ginseng relative to pupae with a preferred reference range of 50-95% ginseng content. Cordycepin, a bioactive nucleoside, increased in the mushroom up to 1.8 fold with the 50% ginseng substitution. Ginseng saponins were present in the fruiting body as well, in their original saponin form and converted into new bioactive saponins, all enriching the final medicinal compound profile of the cordyceps mushrooms.


Reishi and Purple Corn

Patent WO2005067581A2 - Functional substrates for growth of culinary and medicinal mushrooms (filed 2005) demonstrates that reishi mushrooms grown on nonpigmented grains such as rice have undetectable levels of anthocyanins while anthocyanins are present in mushrooms grown on pigmented grain.Anthocyanins are antioxidant plant pigment compounds - red, purple and blue - commonly found in dark berries, fruits and vegetables, and not endogenous to reishi mushrooms. Additionally, heat treatment tends to increase bioavailability of the anthocyanins present in these purple corn-grown mushrooms.


Therapeutic Applications of Anthocyanins

Multiple randomized control trials and meta-analysis with many participants have demonstrated that anthocyanins improve blood lipid profiles, glucose metabolism and endothelial function (Speer 2020). Anthocyanins also improve verbal fluency, memory and brain activity in older adults and may help reduce fatigue, anxiety and depression in younger adults (Sandoval-Ramírez 2022, Ma 2025).


Mushroom

Substrate integration

Composition changes 

Lentinula edodes

Astragalus

↑ Monosaccharide profile shifted toward astragalus-type rhamnose/galacturonic acid 

↑ Total polysaccharides

Pleurotus ostreatus

Chinese formula

↑ Phenolics, flavonoids, terpenoids, and vitamin C

Flammulina velupitas 

Green tea

↑ Caffeine 

Pleurotus ostreatus

Spent coffee grounds

↑ Caffeine, theobromine

Ganoderma lucidum

Ginseng

↑ Ginsenosides and derivatives

Pleurotus ostreatus, Hericium erinaceus, and Flammulina velutipes

Soy

↑ Isoflavones and derivatives (mycelium, unknown if taken up by fruiting body)

Cordyceps sinensis

Ginseng

↑ Ginsenosides

Ganoderma lucidum

Purple corn

↑ Anthocyanins

ree

Influences on Endogenous Medicinal Compounds in Mushrooms

Polysaccharides

Polysaccharide-rich substrate increases polysaccharide yield in mushrooms and the type of bark matters. A 2023 study compared Pleurotus ostreatus and P. eryngii grown on diverse substrates - oat and barley, beech wood shavings, coffee residue and rice bark - against a control of wheat straw. Barley and oat substrate, notably high in polysaccharides, significantly increased polysaccharide yields in the mushrooms (Diamantopoulou 2023).


Another study analyzed Reishi (Ganoderma lucidum) grown on different wood-based substrates (60% wood chips and 40% sawdust by weight). The wood chips were comprised of either birch (Betula spp.), poplar (Populus tremula), spruce (Picea abies), pine (Pinus sylvestrus) or larch (Larix spp.). The highest β-glucan content was found in the mushrooms grown on the poplar wood-based substrate which also had a higher probability of fruiting (Cortina-Escribano 2020).


Therapeutic Applications of Polysaccharides

Mushroom polysaccharides, especially 𝛽-glucans, are often the gold standard for mushroom quality and have a wide range of health benefits. They modulate the immune system via activation of macrophages, natural killer cells, and dendritic cells, facilitating a more effective immune response to pathogens and abnormal cells (Vetvicka 2019). They are also anti-inflammatory, downregulating cytokines like TNF-a and IL-6. Polysaccharides are also antioxidant, scavenging free radicals to protect cells from oxidative stress. Current research has been focusing on polysaccharides as prebiotics - food for beneficial gut microbiota - which improves overall microbiome balance and post biotic production, supporting overall gut health and emphasizing that many of the systemic benefits of polysaccharides may be due to various gut-organ axes (Zhang 2021, Zhao 2023, Wu 2020, Jayachandran 2017, Fernandes 2023).


Ergothioneine 

Nitrogen-rich substrates and LED light exposure increases ergothioneine yield in mushrooms. A 2025 study experimented with Pleurotus ostreatus grown on substrates made from food waste (e.g., kitchen scraps, coffee grounds, barley tea grounds) vs. standard sawdust medium. They found that oyster mushrooms grown on food waste substrates had about twice the ergothioneine content compared to mushrooms grown on sawdust medium. This increase correlated with higher proportions of animal-derived nitrogen in the substrate, suggesting nitrogen impacts ergothioneine biosynthesis. Additionally adding blue LED light during fruiting further enhanced ergothioneine content (Katoaka 2025).


A study investigating the chemistry of lion's mane mushroom analyzed extraction methods and growing conditions, revealing significantly higher ergothioneine content when grown on a substrate of 55% hemp straw and 45% wheat bran, compared to 55% beech sawdust and 45% wheat bran. As expected due to its water solubility, ergothioneine content was also much higher in a dual extract of ethanol and water compared to ethanol alone. Notably, the beech sawdust substrate proved ideal for all other bioactive compounds, including lovastatin, phenolic compounds, and β-glucans (Kala 2025) .


Ergothioneine is synthesized by fungi from histidine and cysteine with synthesis potentially regulated by nitrogen availability. Foods rich in protein/nitrogen, especially animal derived kitchen waste appear to upregulate ergothioneine biosynthesis pathways. Stressors also boost ergothioneine production -  In addition to nitrogen-rich food waste, LED light exposure has shown an increase in ergothioneine content in multiple other studies. 


LED light exposure increased ergothioneine content in shiitake, Lentinula edodes, substantially by 2.8mg/g dry weight vs 1.2mg/6 under white light or control conditions.LED light exposure also shortened the cultivation period and improved antioxidant activity (Kim 2024). 


Light intensities and colors generally impact bioactive compounds in oyster mushrooms. In a 2022 study researchers found that the cap and stem distribution differed, riboflavin increased under higher light, and blue and white LEDs notably improved commercial yield and ergothioneine content compared to low-light controls (Zawadzka 2022).


Therapeutic Applications of Ergothioneine

Ergothioneine is a unique amino acid with significant antioxidant and cytoprotectant properties. The body actively concentrates ergothioneine in tissues, generally more susceptible to oxidative stress, via the OCTN1 transporter where it acts as an antioxidant. It also supports immune and vascular health, lowering vascular inflammation(Zhang 2025, Chen 2025). Human trials have also demonstrated that higher blood levels of ergothioneine are linked to healthier aging, reduced cardiovascular disease, cognitive decline and all-cause mortality. Lower levels of ergothioneine have also been linked in older adults with neurodegenerative disease (Tian 2023).


Cordycepin

Insect-based substrates increase cordycepin content in cordyceps mushrooms and so do certain medicinal plants. Cordyceps is known for its entomopathogenic nature: it is an insect-parasitic fungus. Considering this, it is not surprising that certain bioactive compounds are more present when the substrate contains insects. We can’t depend on this relationship for large scale production - wild-harvested cordyceps are rare and because of that, very expensive and inaccessible to most. Most commercial cordyceps products are cordyceps myceliated grain, typically grown on brown rice and sometimes sorghum, millet or others. A better option, though possibly less scalable, is insect-based substrate. 


When cordyceps was grown on insect-based substrates, cordycepin reached ~2.93 mg/g dry weight and adenosine ~1.06 mg/g, a significant increase compared to cordyceps grown without insects. Brihaspa atrostigmella larvae residues were the top substrate for these increases, outperforming others like cricket/fly residues ( Nguyen 2024). If insect residue is not available, another way to increase cordycepin in mushroom yield is to inhibit cordycepin degradation by including adenosine deaminase (ADA)-inhibiting plants. Researchers included eight medicinal plants to include in brown rice substrate - green tea, turmeric rhizome, linseed, Chinese lizard tail, wild yam rhizome, white mulberry leaves, angelica root and slippery elm bark. The medicinal plants were each combined with brown rice at 15% and 25%. Cordyceps grew on all substrates, but dramatically differently, and contained different amounts of cordycepin with the proposed mechanism being ADA-inhibiting compounds in the plants. Overall, seven samples grown with 25% medicinal plant concentration had a greater yield of cordycepin than those grown on the 15%. The exception was green tea, Camelia sinensis, which had a higher cordycepin content at 15%. The greatest cordycepin concentration was seen on cordyceps grown on white mulberry leaves, Mori folium, at 42.3mg/g dry weight. (Turk 2023). 


Therapeutic Applications of Cordycepin

Cordycepin has demonstrated a number of potential health benefits. Human clinical trials are limited, but in vitro and animal research has demonstrated  anti-inflammatory activity via inhibition of NF-κB signaling and reduced pro-inflammatory cytokines (Tuli 2013). It has also shown antitumor activity - promoting apoptosis, inhibiting tumor cell proliferation and suppressing metastasis - through mTOR inhibition pathways (Das 2010). Cordycepin is also antioxidant and neuroprotective, it has been shown to protect neuronal cells against ischemia and degenerative damage (Nakamura 2014). There may also be metabolic benefits via improved insulin sensitivity and lipid metabolism (Paterson 2008). 


Triterpenes

Wood log cultivation, olive oil and copper and various light exposure increases triterpenes in reishi fruiting bodies. Wood log cultivation (oak, maple, beech and birch logs) compared with substitute cultivation (broadleaf tree sawdust, bran, corn flour, rice malt, gypsum power and other materials) yielded significantly higher triterpenes while there was no significant difference in polysaccharide content (Luo 2024). Olive oil and copper supplementation of sunflower hull based substrates also increased triterpene contents in reishi fruiting bodies (Bidegain 2019).  Light quality and intensity also plays a role in triterpene synthesis. Continuous green light exposure during fruiting increased total triterpenes by 28% in the pileus vs. control (Liu 2024)


Therapeutic Applications of Triterpenes

Triterpenes from medicinal mushrooms have demonstrated antiinflammatory effects, anticancer activity and are hepatoprotective, antioxidant and immune modulating. Their anti-inflammatory effects are through NF-κB inhibition and they reduce pro-inflammatory cytokine levels. Anticancer activity has been observed via apoptosis, antiangiogenesis and antimetastatic effects in preclinical studies (Boh 2007, baby 2017, Dasgupta 2019).


Hericenones and Erinacines

Olive mill solid waste and darkness increases hericenones and erinacines in lion’s mane mushrooms. Olive mill solid waste, a byproduct of olive oil processing, was added to a substrate of eucalyptus sawdust and malt waste at concentrations ranging from 0% to 80%. Using advanced metabolomic profiling, researchers found that olive mill solid waste significantly increased the content of neuroactive hericenones in the fruiting bodies and erinacines in the mycelium (Khatib 2025). Erinacines content is also influenced by the substrate it is colonizing. Turk et. al found that the combination of sawdust 42% + cottonseed meal 12% + rice bran 7% + cottonseed hulls 19% + beet 13% increased erinacine A concentration in lion’s mane mycelium from 1.93 to 3.46mg/g dry weight. They also observed that dark conditions were more optimal than light conditions for erinacine production (Turk 2021, Kostanda 2024).  


Therapeutic Applications of Hericenones and Erinacines

Hericenones and erinacines are bioactive compounds primarily found in the lion's mane fruiting body and mycelium respectively. In laboratory studies, they have been shown to stimulate nerve growth factor which helps to support neuronal health and neuroplasticity. Although hericenones stimulate nerve growth factor synthesis in vitro, their effect is less significant that that of erinacines, which repeatedly demonstrate more significant increases in nerve growth factor activity (Li 2018, Spangenberg 2025, Pangenberg 2025).


Psilobycin 

The addition of peat-vermiculite casings, gypsum and tryptophan derivatives increase psilocybin in Psylocybe mushrooms. The content of psilocybin, the primary psychoactive compound in Psylocybe spp., varies greatly across strains, even when grown under similar conditions (Goff 2024). Research exploring substrate influence on specific strains is limited, but there are a few studies. A 2023 experiment with Psilocybe cubensis found that small changes in the growth matrix, using a peat-vermiculite casing and adding gypsum to popcorn grain and horse manure, produced significant increases in psilocybin content (Foster 2023). A 2023 review also noted that supplementing substrates with tryptophan derivatives can increase psilocybin concentrations in fruiting bodies (Luz 2025).


Therapeutic Applications of Psilocybin

There have been many clinical trials demonstrating that psilocybin can facilitate rapid and sustained reductions in depression. Randomized clinical trials with patients who have major depressive disorder show significant improvement (Davis 2021). Studies have also shown that a single high dose psilocybin session leads to improvements in mood and quality of life in cancer patients with anxiety and depression (Griffiths 2016). Psilocybin may also influence cognitive flexibility and emotional regulation suggesting applications for addiction treatment and other mental health issues. Smaller clinical trials have also shown improvement in cluster headaches, with some individuals reporting full remission (Schindler 2015).


Bioactive Compounds

Mushrooms

Influential Conditions

Polysaccharides

Pleurotus ostreatus and P. eryngii

Ganoderma lucidum

Barley and oat substrate


Poplar wood chips (60% wood chips and 40% sawdust by weight)

Ergothioneine

Pleurotus ostreatus


Lentinula edodes


Hericium erinaceus

Nitrogen-rich food waste

Blue LED Light


LED light


55% hemp straw and 45% wheat bran

Cordycepin

Cordyceps militaris

Insect-based substrate - Brihaspa atrostigmella larvae residues 


15% white mulberry leaves

Triterpenes

Ganoderm lucidum

Wood log cultivation

Olive oil + Copper

Continuous green light

Hericenones and Erinacines


Erinacines

Hericium erinaceus

Olive mill solid waste


Darkness

Psilocybin

Psilocybe cubensis

Tryptophan derivatives

Peat-vermiculite casing and gypsum

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Conclusions

When it comes to bioactive compounds present in culinary and medicinal mushrooms, substrate matters. In the wild, variations in environmental stressors and host trees influence synthesis of endogenous secondary metabolites and in some cases, tree-specific compounds. For this reason, wild mushrooms are preferred by some, though with the understanding that wild mushrooms have more opportunity to contain heavy metals and toxins from an uncontrolled environment.


Research exploring specific tree-derived compounds in medicinal mushrooms is lacking, but a lot can be learned from studies exploring the presence of bioactive plant-derived compounds in mushrooms grown on medicinal plant-integrated substrates. Immunomodulating activity of shiitake becomes more potent when grown on astragalus; oyster mushrooms contain more antioxidant secondary metabolites when grown on a TCM formulas; enoki and oyster mushrooms take up caffeine from green tea and spent coffee grounds; reishi takes up and biotransforms potent ginsenosides from ginseng; mushrooms grown on soy-based substrate likely contain estrogen-modulating isoflavones; cordyceps grown on ginseng synthesizes more cordycepin while also taking up and transforming bioactive saponins; and reishi grown on purple corn contains antioxidant anthocyanins. Influencing the chemical makeup of medicinal mushrooms to contain specific bioactive plant constituents is the future of medicinal mushroom cultivation.


As exemplified, there are many ways to enrich the already therapeutic chemistry of mushrooms with plant-derived compounds. There are also numerous ways to enhance the endogenous medicinal compounds of mushrooms. Polysaccharide-rich substrate and specific bark profiles influence polysaccharide content; nitrogen-rich substrates, hemp and LED light exposure increases ergothioneine yield; insect-based substrates and various medicinal plant based substrates increase cordycepin content in cordyceps; wood log cultivation, olive oil and copper and continuous green light exposure increases triterpenes in reishi fruiting bodies; olive waste increases hericenones and erinacines in lion’s mane; and tryptophan increases psilocybin in Psilocybe mushrooms.


Currently, the most comprehensive labeling on medicinal mushroom products specifies the names of the mushrooms in the formula, their concentration, and perhaps the amount of 𝛽-glucans, triterpenes and ergothioneine present. In the case of myceliated grain products, the substrate is occasionally listed as an “other ingredient”. In the future, I anticipate the inclusion of more detailed information about what the mushroom was growing on - type of host tree, bark, or logs, and a more thorough description of the substrate's composition, and at least noting any medicinal plant additives. Perhaps the future of mushroom coffee products will feature mushrooms cultivated on spent coffee grounds, and adaptogenic tea blends may incorporate not only ginseng, but also reishi and cordyceps grown on ginseng-enriched substrate containing their distinctive mushroom-transformed ginsenosides. Until then, I’ll continue to mix my herbs and mushrooms in formula and use both wild and cultivated mushrooms as appropriate.  











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