Ursolic acid

Ursolic acid at a Glance

Ursolic acid is a pentacyclic triterpenoid concentrated in the waxy cuticle of apple peel and in culinary Lamiaceae herbs such as rosemary, sage and thyme — a structural component of the plant’s surface rather than an internal metabolite.

Its marquee signal is skeletal-muscle preservation, and the way it was found is unusual: it emerged from a gene-signature screen looking for compounds that reverse the transcriptional pattern of muscle atrophy, rather than from traditional use or a mechanistic hypothesis.

Its other research lines — anti-inflammatory, immune and anticancer — are broader and more conventional.

What It's Used For

Ursolic acid is a pentacyclic triterpenoid of the ursane class — the waxy, lipophilic acid that waterproofs the cuticle of apple peel and accumulates in the leaves of rosemary, sage, thyme, bearberry, holy basil, lemon balm, catnip and many other plants here. It carries one of the broadest preclinical literatures of any single triterpene: isolated ursolic acid shows skeletal-muscle-preserving, anti-inflammatory, anticancer, metabolic and GABAergic activity across cell and rodent models ^{7,12Reference 7Checker R et al. · 2012In vitroPotent anti-inflammatory activity of ursolic acid, a triterpenoid antioxidant, is mediated through suppression of NF-κB, AP-1 and NF-AT — in vitro / ex vivoView study →Reference 12Sun Q et al. · 2020Systematic reviewUrsolic acid: a systematic review of its pharmacology, toxicity and rethink on its pharmacokinetics based on PK-PD model — reviewView study →}.

Two facts frame all of it. First, almost every result is preclinical or in vitro — the isolated-molecule human data are limited to a few small, mixed muscle pilots. Second, and more decisive, ursolic acid is a poorly water-soluble BCS class IV compound: oral bioavailability is very low, plasma half-life is under an hour, and it is a substrate for efflux and CYP metabolism ^{11,12Reference 11Zhu Z et al. · 2013Clinical trialA phase I pharmacokinetic study of ursolic acid nanoliposomes in healthy volunteers and patients with advanced solid tumors — phase I clinical trialView study →Reference 12Sun Q et al. · 2020Systematic reviewUrsolic acid: a systematic review of its pharmacology, toxicity and rethink on its pharmacokinetics based on PK-PD model — reviewView study →}.

Concentrations that work in a dish are not trivially reached from an oral dose, so the potency seen in vitro should be read with heavy discount. Within the source herbs it is usually a supporting constituent in a matrix — hawthorn’s own literature notes that isolated triterpene acids show little of the whole extract’s cardiovascular effect ^{10Reference 10Occhiuto F et al. · 1986Comparative study of the cardiovascular activity of shoots, leaves and flowers of Crataegus oxyacantha: 2}.

Evidence by IndicationStrength of support · Beta

Support is an experimental score — weighted by study type (human > animal > in vitro) and volume, tempered for isolate-vs-plant match and bioavailability. A beta, not a validated metric. Each application links to its write-up.

01Skeletal muscle & atrophyPromising46%Isolated UA; deep mouse mechanism (Akt/IGF-I) + small, mixed human pilots ^{1,3,4Reference 1Kunkel SD et al. · 2011AnimalmRNA expression signatures of human skeletal muscle atrophy identify a natural compound that increases muscle mass — Connectivity Map screen + mouse in vivoView study →Reference 3Bang HS et al. · 2017RCTUrsolic acid supplementation decreases markers of skeletal muscle damage during resistance training in resistance-trained men — randomised pilot trial (450 mg/day, 8 weeks)View study →Reference 4Lobo PCB et al. · 2021RCTUrsolic acid has no additional effect on muscle strength and mass in active men undergoing a high-protein diet and resistance training — double-blind placebo-controlled trialView study →}

02Anti-inflammatory & immuneShaky38%Isolated UA; resolved NF-κB/AP-1/NF-AT mechanism, in-vitro/ex-vivo ^{7Reference 7Checker R et al. · 2012In vitroPotent anti-inflammatory activity of ursolic acid, a triterpenoid antioxidant, is mediated through suppression of NF-κB, AP-1 and NF-AT — in vitro / ex vivoView study →}

03AnticancerShaky34%Isolated UA; broad cultured-tumor-line data, high doses; sparse in vivo ^{12Reference 12Sun Q et al. · 2020Systematic reviewUrsolic acid: a systematic review of its pharmacology, toxicity and rethink on its pharmacokinetics based on PK-PD model — reviewView study →}

04Metabolic (glucose, obesity, lipids)Shaky30%Isolated UA; diet-induced-obesity mouse model; no isolate human data ^{2Reference 2Kunkel SD et al. · 2012AnimalUrsolic acid increases skeletal muscle and brown fat and decreases diet-induced obesity, glucose intolerance and fatty liver disease — mouse in vivo animal modelView study →}

05Anxiolytic (GABA-transaminase)Shaky26%Isolated UA in vitro (GABA-T inhibition); human evidence is whole lemon-balm extract ^{8,9Reference 8Awad R et al. · 2009In vitroBioassay-guided fractionation of lemon balm (Melissa officinalis L.) using an in vitro measure of GABA-transaminase activity — in vitroView study →Reference 9Ibarra A et al. · 2010AnimalEffects of chronic administration of Melissa officinalis L. extract on anxiety-like reactivity and on circadian and exploratory activities in mice — mouse modelView study →}

06CardiovascularShaky22%One isolated-principle rabbit-heart study; whole-hawthorn matrix caveat ^{10Reference 10Occhiuto F et al. · 1986Comparative study of the cardiovascular activity of shoots, leaves and flowers of Crataegus oxyacantha: 2}

Mechanisms

Pharmacokinetics

Bioavailability is the load-bearing caveat for the whole ursolic-acid literature. It is a highly lipophilic, poorly water-soluble pentacyclic triterpene — a BCS class IV molecule (low solubility and low permeability) — so its oral bioavailability as the pure compound is very low, the result of poor dissolution, extensive pre-systemic metabolism (including by CYP3A4) and efflux ^{11,12Reference 11Zhu Z et al. · 2013Clinical trialA phase I pharmacokinetic study of ursolic acid nanoliposomes in healthy volunteers and patients with advanced solid tumors — phase I clinical trialView study →Reference 12Sun Q et al. · 2020Systematic reviewUrsolic acid: a systematic review of its pharmacology, toxicity and rethink on its pharmacokinetics based on PK-PD model — reviewView study →}. In rodents it is absorbed quickly (Tmax ~30 min) but cleared quickly too, with a plasma half-life under about an hour ^{12Reference 12Sun Q et al. · 2020Systematic reviewUrsolic acid: a systematic review of its pharmacology, toxicity and rethink on its pharmacokinetics based on PK-PD model — reviewView study →}. The one human pharmacokinetic study used a nanoliposome formulation given intravenously to healthy volunteers and cancer patients precisely because plain oral ursolic acid does not achieve useful plasma levels; exposure rose roughly dose-proportionally and the formulation was tolerated up to defined limits ^{11Reference 11Zhu Z et al. · 2013Clinical trialA phase I pharmacokinetic study of ursolic acid nanoliposomes in healthy volunteers and patients with advanced solid tumors — phase I clinical trialView study →}. The practical upshot mirrors osthole and berberine: the potent in-vitro concentrations are not reachable from eating apple peel or taking an unformulated capsule, which tempers every downstream claim and is why the applied literature leans on nanoparticles, liposomes, phytosomes and bioenhancers (e.g. piperine) ^{11,12Reference 11Zhu Z et al. · 2013Clinical trialA phase I pharmacokinetic study of ursolic acid nanoliposomes in healthy volunteers and patients with advanced solid tumors — phase I clinical trialView study →Reference 12Sun Q et al. · 2020Systematic reviewUrsolic acid: a systematic review of its pharmacology, toxicity and rethink on its pharmacokinetics based on PK-PD model — reviewView study →}.

Clinical trials

There is no large trial of isolated ursolic acid for any indication. The human evidence is three small supplementation studies in the muscle space — a positive pilot on damage markers ^{3Reference 3Bang HS et al. · 2017RCTUrsolic acid supplementation decreases markers of skeletal muscle damage during resistance training in resistance-trained men — randomised pilot trial (450 mg/day, 8 weeks)View study →}, a negative controlled trial on strength/mass ^{4Reference 4Lobo PCB et al. · 2021RCTUrsolic acid has no additional effect on muscle strength and mass in active men undergoing a high-protein diet and resistance training — double-blind placebo-controlled trialView study →}, and a negative loquat-leaf (Ua-standardized) trial ^{5Reference 5Cho YH et al. · 2016RCTEffect of loquat leaf extract on muscle strength, muscle mass, and muscle function in healthy adults — randomised, double-blind, placebo-controlled trial (~51 mg ursolic acid/day)View study →} — plus an early-phase nanoliposome pharmacokinetic/safety study in volunteers and cancer patients ^{11Reference 11Zhu Z et al. · 2013Clinical trialA phase I pharmacokinetic study of ursolic acid nanoliposomes in healthy volunteers and patients with advanced solid tumors — phase I clinical trialView study →}. Everything else is preclinical.

Last checked: July 2026.

Isolate vs. Plant Studies

Ursolic acid sits at both ends of the isolate-vs-plant spectrum. Its marquee muscle, metabolic, anti-inflammatory and anticancer findings all use the isolated molecule (or defined synthetic/nanoparticle preparations), so those describe the compound rather than a plant standing in for it ^{1,2,7,12Reference 1Kunkel SD et al. · 2011AnimalmRNA expression signatures of human skeletal muscle atrophy identify a natural compound that increases muscle mass — Connectivity Map screen + mouse in vivoView study →Reference 2Kunkel SD et al. · 2012AnimalUrsolic acid increases skeletal muscle and brown fat and decreases diet-induced obesity, glucose intolerance and fatty liver disease — mouse in vivo animal modelView study →Reference 7Checker R et al. · 2012In vitroPotent anti-inflammatory activity of ursolic acid, a triterpenoid antioxidant, is mediated through suppression of NF-κB, AP-1 and NF-AT — in vitro / ex vivoView study →Reference 12Sun Q et al. · 2020Systematic reviewUrsolic acid: a systematic review of its pharmacology, toxicity and rethink on its pharmacokinetics based on PK-PD model — reviewView study →}. But almost all of that is preclinical, and the small human muscle trials are mixed-to-negative ^{3,4,5Reference 3Bang HS et al. · 2017RCTUrsolic acid supplementation decreases markers of skeletal muscle damage during resistance training in resistance-trained men — randomised pilot trial (450 mg/day, 8 weeks)View study →Reference 4Lobo PCB et al. · 2021RCTUrsolic acid has no additional effect on muscle strength and mass in active men undergoing a high-protein diet and resistance training — double-blind placebo-controlled trialView study →Reference 5Cho YH et al. · 2016RCTEffect of loquat leaf extract on muscle strength, muscle mass, and muscle function in healthy adults — randomised, double-blind, placebo-controlled trial (~51 mg ursolic acid/day)View study →}. The source-herb roles run the other way: within hawthorn, bearberry, lemon balm and yerba maté, ursolic acid is a supporting triterpene in a matrix, and the human clinical effects of those herbs (hawthorn’s cardiotonic action, lemon balm’s anxiolysis) are whole-plant results that cannot be reassigned to ursolic acid — hawthorn’s literature explicitly finds isolated constituents underperform the whole extract ^{10Reference 10Occhiuto F et al. · 1986Comparative study of the cardiovascular activity of shoots, leaves and flowers of Crataegus oxyacantha: 2}. Read the preclinical work as evidence for the molecule and the herb trials as evidence for the plants.

Prevalence in Nature

Ursolic acid is one of the most widely distributed plant triterpenoids, concentrated in the epicuticular wax of fruit peels and in the leaves and aerial parts of many families — most prominently the mint family (Lamiaceae) and the heath, rose, apple, myrtle and holly families ^{12,13Reference 12Sun Q et al. · 2020Systematic reviewUrsolic acid: a systematic review of its pharmacology, toxicity and rethink on its pharmacokinetics based on PK-PD model — reviewView study →Reference 13Jäger S et al. · 2009Pentacyclic triterpene distribution in various plants — rich sources for a new group of multi-potent plant extracts — analytical surveyView study →}. It occurs chiefly in surface tissue: fruit cuticle, leaf epidermis and flowers, rather than roots. By measured concentration the richest sources are culinary Lamiaceae leaves — rosemary at ~2.95% dry weight, sage ~1.8%, lavender ~1.6% and thyme ~0.9% — with bearberry leaf ~1.2% ^{13Reference 13Jäger S et al. · 2009Pentacyclic triterpene distribution in various plants — rich sources for a new group of multi-potent plant extracts — analytical surveyView study →}. Apple peel, the compound’s iconic dietary source, is actually lower on a concentration basis (~0.33–0.58% of dry weight, i.e. ~3–6 mg/g), though because the peel of one medium apple holds roughly 40–50 mg it is a meaningful everyday exposure ^{14Reference 14Butkevičiūtė A et al. · 2022Triterpene content in flesh and peel of apples grown on different rootstocks — analytical studyView study →}. It co-occurs almost universally with its regioisomer oleanolic acid (oleanane skeleton) at lower levels, the two differing only in the position of one methyl group.

Biosynthetically it is a triterpenoid built from the universal C30 precursor 2,3-oxidosqualene: an oxidosqualene cyclase (α-amyrin synthase) folds the chain into the pentacyclic ursane skeleton α-amyrin, which cytochrome-P450 enzymes of the CYP716 family then oxidize stepwize at C-28 to the carboxylic acid, yielding ursolic acid (the parallel oleanane route gives oleanolic acid) ^{16Reference 16Yasumoto S et al. · 2017Functional characterization of CYP716 family P450 enzymes in triterpenoid biosynthesis in tomato — plant biochemistryView study →}. It is essentially a plant product; there is no established endogenous animal source.

Discovery & Synthesis

Ursolic acid was identified in the epicuticular wax of apples in the 1920s, among the first triterpenoids characterized from a fruit cuticle; its name derives from this waxy origin, and it carries the older synonyms urson, prunol and malol (the last two reflecting early isolation from Prunus and Malus fruit) ^{12Reference 12Sun Q et al. · 2020Systematic reviewUrsolic acid: a systematic review of its pharmacology, toxicity and rethink on its pharmacokinetics based on PK-PD model — reviewView study →}. Its molecular formula is C₃₀H₄₈O₃ and its systematic name is 3β-hydroxyurs-12-en-28-oic acid. A practical, scalable total chemical synthesis is not well documented — the pentacyclic ursane skeleton is synthetically demanding, and there is no commercial synthetic route. Consequently ursolic acid is manufactured entirely by extraction from plant material: apple pomace (a cheap by-product of juice and cider pressing), rosemary and sage leaf, and Eriobotrya (loquat) leaf are the usual feedstocks, purified by solvent extraction and chromatography ^{13,14Reference 13Jäger S et al. · 2009Pentacyclic triterpene distribution in various plants — rich sources for a new group of multi-potent plant extracts — analytical surveyView study →Reference 14Butkevičiūtė A et al. · 2022Triterpene content in flesh and peel of apples grown on different rootstocks — analytical studyView study →}. Most of the medicinal-chemistry effort has gone not into synthesizing the parent molecule but into semi-synthetic C-3/C-28 derivatives and into formulation (nanoparticles, liposomes) to overcome its poor bioavailability ^{11,12Reference 11Zhu Z et al. · 2013Clinical trialA phase I pharmacokinetic study of ursolic acid nanoliposomes in healthy volunteers and patients with advanced solid tumors — phase I clinical trialView study →Reference 12Sun Q et al. · 2020Systematic reviewUrsolic acid: a systematic review of its pharmacology, toxicity and rethink on its pharmacokinetics based on PK-PD model — reviewView study →}.

Patents: not yet researched (future patent-loop pass).

Toxicity & Safety

Ursolic acid carries a low toxicity flag. It is a normal component of the human diet — present in the skins of apples and many fruits and in common culinary herbs — and reviews describe it as generally well tolerated in animal studies, with no signature acute organ toxicity at the doses used and a wide preclinical safety margin ^{12Reference 12Sun Q et al. · 2020Systematic reviewUrsolic acid: a systematic review of its pharmacology, toxicity and rethink on its pharmacokinetics based on PK-PD model — reviewView study →}. The small human muscle pilots reported no serious adverse events at 450 mg/day for 8–12 weeks ^{3,4Reference 3Bang HS et al. · 2017RCTUrsolic acid supplementation decreases markers of skeletal muscle damage during resistance training in resistance-trained men — randomised pilot trial (450 mg/day, 8 weeks)View study →Reference 4Lobo PCB et al. · 2021RCTUrsolic acid has no additional effect on muscle strength and mass in active men undergoing a high-protein diet and resistance training — double-blind placebo-controlled trialView study →}. The main theoretical concern is metabolic: ursolic acid is a substrate for and inhibitor of CYP3A4, and formulation work deliberately pairs it with the CYP inhibitor piperine to raise exposure, so at high supplemental intakes an interaction with CYP3A4-cleared medications is plausible though undocumented in humans ^{11,12Reference 11Zhu Z et al. · 2013Clinical trialA phase I pharmacokinetic study of ursolic acid nanoliposomes in healthy volunteers and patients with advanced solid tumors — phase I clinical trialView study →Reference 12Sun Q et al. · 2020Systematic reviewUrsolic acid: a systematic review of its pharmacology, toxicity and rethink on its pharmacokinetics based on PK-PD model — reviewView study →}. A formal human safety database and interaction studies do not yet exist because the isolated molecule has not been trialled at scale, so absence of reported harm should not be read as established safety. As with all constituents on this database, the herbs that supply ursolic acid carry their own whole-plant cautions (for example hawthorn’s potential to add to cardiac-glycoside drugs) that are separate from any property of ursolic acid itself.

Dosage

There is no established human dose for isolated ursolic acid. The small muscle pilots used 450 mg per day of the isolated compound over 8 weeks ^{3,4Reference 3Bang HS et al. · 2017RCTUrsolic acid supplementation decreases markers of skeletal muscle damage during resistance training in resistance-trained men — randomised pilot trial (450 mg/day, 8 weeks)View study →Reference 4Lobo PCB et al. · 2021RCTUrsolic acid has no additional effect on muscle strength and mass in active men undergoing a high-protein diet and resistance training — double-blind placebo-controlled trialView study →}; a loquat-leaf trial delivered ~51 mg/day of ursolic acid over 12 weeks ^{5Reference 5Cho YH et al. · 2016RCTEffect of loquat leaf extract on muscle strength, muscle mass, and muscle function in healthy adults — randomised, double-blind, placebo-controlled trial (~51 mg ursolic acid/day)View study →}; and preclinical rodent work commonly uses tens to a couple of hundred milligrams per kilogram, figures inflated to compensate for poor absorption ^{1,2Reference 1Kunkel SD et al. · 2011AnimalmRNA expression signatures of human skeletal muscle atrophy identify a natural compound that increases muscle mass — Connectivity Map screen + mouse in vivoView study →Reference 2Kunkel SD et al. · 2012AnimalUrsolic acid increases skeletal muscle and brown fat and decreases diet-induced obesity, glucose intolerance and fatty liver disease — mouse in vivo animal modelView study →}.

Because oral bioavailability is very low and metabolism rapid, plain oral ursolic acid reaches only low plasma levels, and bioavailability-enhanced forms (nanoparticles, liposomes, piperine-paired) change exposure substantially ^{11,12Reference 11Zhu Z et al. · 2013Clinical trialA phase I pharmacokinetic study of ursolic acid nanoliposomes in healthy volunteers and patients with advanced solid tumors — phase I clinical trialView study →Reference 12Sun Q et al. · 2020Systematic reviewUrsolic acid: a systematic review of its pharmacology, toxicity and rethink on its pharmacokinetics based on PK-PD model — reviewView study →}. These are doses studied in research and are not a personal recommendation.

References

1. Kunkel SD, Suneja M, Ebert SM, Bongers KS, Fox DK, Malmberg SE, et al. (2011). mRNA expression signatures of human skeletal muscle atrophy identify a natural compound that increases muscle mass — Connectivity Map screen + mouse in vivo. Cell Metabolism, 13(6), 627–638. https://pubmed.ncbi.nlm.nih.gov/21641545/
2. Kunkel SD, Elmore CJ, Bongers KS, Ebert SM, Fox DK, Dyle MC, et al. (2012). Ursolic acid increases skeletal muscle and brown fat and decreases diet-induced obesity, glucose intolerance and fatty liver disease — mouse in vivo animal model. PLoS ONE, 7(6), e39332. https://pubmed.ncbi.nlm.nih.gov/22745735/
3. Bang HS, Seo DY, Chung YM, Kim DH, Lee SJ, Lee SR, et al. (2017). Ursolic acid supplementation decreases markers of skeletal muscle damage during resistance training in resistance-trained men — randomised pilot trial (450 mg/day, 8 weeks). Korean Journal of Physiology & Pharmacology, 21(6), 651–656. https://pubmed.ncbi.nlm.nih.gov/29200908/
4. Lobo PCB, Vieira IP, Pichard C, Marques ADS, Pimentel GD. (2021). Ursolic acid has no additional effect on muscle strength and mass in active men undergoing a high-protein diet and resistance training — double-blind placebo-controlled trial. Clinical Nutrition, 40(2), 581–589. https://pubmed.ncbi.nlm.nih.gov/32593524/
5. Cho YH, Lee SY, Kim CM, Kim ND, Choe S, et al. (2016). Effect of loquat leaf extract on muscle strength, muscle mass, and muscle function in healthy adults — randomised, double-blind, placebo-controlled trial (~51 mg ursolic acid/day). Evidence-Based Complementary and Alternative Medicine, 2016, 4301621. https://pubmed.ncbi.nlm.nih.gov/27999607/
6. Fernandes ES, Machado Mde O, Becker AM, de Andrade F, Maraschin M, da Silva EL. (2012). Yerba maté (Ilex paraguariensis) enhances the gene modulation and activity of paraoxonase-2 — in vitro and in vivo studies (whole-extract, ursolic acid among candidate actives). Nutrition, 28(11–12), 1157–1164. https://pubmed.ncbi.nlm.nih.gov/22964087/
7. Checker R, Sandur SK, Sharma D, Patwardhan RS, Jayakumar S, Kohli V, et al. (2012). Potent anti-inflammatory activity of ursolic acid, a triterpenoid antioxidant, is mediated through suppression of NF-κB, AP-1 and NF-AT — in vitro / ex vivo. PLoS ONE, 7(2), e31318. https://pubmed.ncbi.nlm.nih.gov/22363615/
8. Awad R, Muhammad A, Durst T, Trudeau VL, Arnason JT. (2009). Bioassay-guided fractionation of lemon balm (Melissa officinalis L.) using an in vitro measure of GABA-transaminase activity — in vitro. Phytotherapy Research, 23(8), 1075–1081. https://pubmed.ncbi.nlm.nih.gov/19165747/
9. Ibarra A, Feuillere N, Roller M, Lesburgere E, Beracochea D. (2010). Effects of chronic administration of Melissa officinalis L. extract on anxiety-like reactivity and on circadian and exploratory activities in mice — mouse model. Phytomedicine, 17(6), 397–403. https://pubmed.ncbi.nlm.nih.gov/20171069/
10. Occhiuto F, Circosta C, Costa R, Briguglio F, Tommasini A. (1986). Comparative study of the cardiovascular activity of shoots, leaves and flowers of Crataegus oxyacantha: 2. Action of extracts and isolated pure active principles on the isolated rabbit heart — isolated-organ study. Plantes Médicinales et Phytothérapie, 20, 52–63.
11. Zhu Z, Qian Z, Yan Z, Zhao C, Wang H, Ying G. (2013). A phase I pharmacokinetic study of ursolic acid nanoliposomes in healthy volunteers and patients with advanced solid tumors — phase I clinical trial. International Journal of Nanomedicine, 8, 129–136. https://pubmed.ncbi.nlm.nih.gov/23319864/
12. Sun Q, He M, Zhang M, Zeng S, Chen L, Zhou L, Xu H. (2020). Ursolic acid: a systematic review of its pharmacology, toxicity and rethink on its pharmacokinetics based on PK-PD model — review. Fitoterapia, 147, 104735. https://pubmed.ncbi.nlm.nih.gov/33010369/
13. Jäger S, Trojan H, Kopp T, Laszczyk MN, Scheffler A. (2009). Pentacyclic triterpene distribution in various plants — rich sources for a new group of multi-potent plant extracts — analytical survey. Molecules, 14(6), 2016–2031. https://pubmed.ncbi.nlm.nih.gov/19513002/
14. Butkevičiūtė A, Janulis V, Kviklys D. (2022). Triterpene content in flesh and peel of apples grown on different rootstocks — analytical study. Plants, 11(9), 1247. https://pubmed.ncbi.nlm.nih.gov/35567248/
15. Micota B, Sadowska B, Podsędek A, Redzynia M, et al. (2014). Leonurus cardiaca L. herb — a derived extract and ursolic acid as factors affecting the adhesion capacity of Staphylococcus aureus in the context of infective endocarditis — in vitro. Acta Biochimica Polonica, 61(2), 385–388. https://pubmed.ncbi.nlm.nih.gov/24918490/
16. Yasumoto S, Seki H, Shimizu Y, Fukushima EO, Muranaka T. (2017). Functional characterization of CYP716 family P450 enzymes in triterpenoid biosynthesis in tomato — plant biochemistry. Frontiers in Plant Science, 8, 21. https://pubmed.ncbi.nlm.nih.gov/28194155/

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