Pharmacognosy · Cardiovascular & Metabolic · Antihypertensive

Olive Leaf

Olea europaea L. — the Mediterranean olive tree, domesticated in Crete around 3500 BC — whose leaf is studied for antihypertensive, cardioprotective, antioxidant, and antidiabetic effects centered on the compound oleuropein.

41 Primary Refs
22 Properties
Leaf Parts Used
✦ Researched
Last Updated
Primary Source Wikiphyto · NCBI PubMed · Phytomedicine
Family Oleaceae
Cardiovascular & Metabolic Support

Biological Overview

The olive is a tree commonly cultivated across Mediterranean regions, reaching about 10 meters in height, with a gray, gnarled and fissured trunk and numerous branches bearing leathery, lance-shaped, grayish-green evergreen leaves. It is a vigorous, long-lived tree with very dense, homogeneous wood.

Life CycleLong-lived evergreen tree
HabitatMediterranean basin
Secoiridoid Content (Leaf)6–14% dry weight
Marker CompoundOleuropein
Botanical Classification

Taxonomy & Identification

Latin Name
Olea europaea L.
Family
Oleaceae
Common Names
Olive, Olive Leaf
Wild Form
Oleaster (spontaneous Mediterranean form)
Parts Used
Leaf, bud, fruit (olive)
Habitat
Native to Syria; naturalized Mediterranean basin
Ethnobotany & Symbolism

History & Tradition

The olive tree is thought to have been domesticated in Crete around 3500 BC. Few trees carry as much symbolic weight: the olive branch is a universal symbol of peace, and in ancient Greece, a crown of olive leaves was placed on the heads of victors at the Olympic Games.

The tree's resilience matches its symbolism — it can live for centuries, tolerates drought and heat, and its main limiting factors are northern cold and extreme aridity, though its cultivable range has extended as far as California, South Africa, and Australia.

⚠ Leaf, Not Fruit

Oleuropein, the compound behind most of this monograph, is barely present in the olive fruit or its oil. The medicinal research summarized here is almost entirely about the leaf — a genuinely different part of the plant from the one used in cooking.

Timeline

~3500 BC

Domestication in Crete

The olive tree is domesticated, beginning millennia of Mediterranean cultivation.

1992

Hypoglycemic Activity Confirmed

Gonzalez et al. document hypoglycemic activity of olive leaf extracts.[22]

2011

Captopril Comparison Trial

Susalit et al. publish a randomized trial comparing standardized olive leaf extract to captopril in stage-1 hypertension.[4]

2017

EMA Assessment

The European Medicines Agency publishes its herbal assessment report on olive leaf.[32]

The Key Molecule

Oleuropein — Deep Dive

The secoiridoid compound behind nearly every property documented for olive leaf — and the standard by which extracts are measured.

🍃

Dominant Leaf Compound

Oleuropein and related secoiridoids make up 6 to 14% of the leaf's dry weight, with some cultivars containing even more.[1]

🎯

ACE-Inhibiting Secoiridoid

Oleuropein, ligustroside, and related secoiridoids inhibit angiotensin-converting enzyme (ACE), directly relevant to blood pressure regulation.[6]

🫀

Superior to Isolated Compounds

An ethanolic leaf extract containing oleuropein alongside hydroxytyrosol, verbascoside, luteolin, and quercetin protects cardiomyocyte viability more effectively than any single one of those compounds alone.[13]

⚗️

Minimal Drug-Metabolism Interference

Oleuropein shows only weak inhibition of the CYP1A2 liver enzyme, suggesting a low risk of pharmacokinetic drug interactions.[36]

⚠ Insulin-Related Mechanism Worth Knowing

Oleuropeoside lowers blood glucose through two distinct mechanisms.

At a dose of 16 mg/kg in animal research, oleuropeoside potentiated glucose-induced insulin release and increased peripheral glucose uptake. This dual mechanism is part of why a theoretical interaction with diabetes medication is worth discussing with a healthcare provider — see Safety & Precautions.[22]

Available Forms

Parts Used & Available Forms

The dried leaf is the primary medicinal part, available in five documented galenic forms.

Mother Tincture

An alcoholic tincture of the leafy branch.

Liquid · Traditional

Glycerin Macerate (Gemmotherapy)

A glycerin macerate of the young shoot, used in the gemmotherapy tradition.

Bud Extract

EPS (Standardized Plant Extract)

A standardized fresh-plant extract of the leaf.

Leaf Extract

Dry Extract

A concentrated dry extract, typically in capsule form — the form used in most clinical trials.

Standardized Capsule

Fluid Extract

A concentrated liquid extract of the leaf.

Liquid Extract

Clinical Dosimetry

Dosages

Four forms, each with its own usual dose as given in the primary source.

Form Dose Frequency
Leaf Decoction 30 g/L 1 liter per day, in discontinuous cures
Glycerin Macerate (Bud), 1°D 50 drops Three times daily, or 1 teaspoon in the morning
EPS (Leaf) 15 mL (3 teaspoons) In 1 liter water, drunk through the day, 10 days/month; or 1 teaspoon each morning
Dry Extract 400–500 mg per capsule Morning and evening (800 mg–1 g/day)
Phytochemistry

Composition

A secoiridoid-rich leaf profile, distinct from the fatty-acid-dominant fruit and oil.

Leaf — Secoiridoids & Triterpenes

Oleuropeoside (Oleuropein)Principal secoiridoid; 6–14% of leaf dry weight
Major
Ligustroside & OleurosideRelated secoiridoids
Present
Oleanolic, Ursolic & Maslinic AcidsTetra- and pentacyclic triterpenes[2]
Present

Leaf — Flavonoids & Phenolics

Luteolin & Apigenin GlycosidesFlavones
Present
Tyrosol & HydroxytyrosolPhenolic derivatives
Present
Verbascoside & CatechinPhenylpropanoid and flavan-3-ol
Present

Fruit — Fatty Oil

Oleic Acid (Omega-9)63–81% of the oil
Major
Linoleic Acid (Omega-6)~15% of the oil
Present
Oleocanthal & TocopherolsVitamin E ~1.6 mg per tablespoon
Present

Plant Properties — Pharmacodynamics

22 properties documented, the richest property profile of any monograph in this series.

22 Properties Antihypertensive Cardioprotective Antidiabetic
🩺

Antihypertensive (Modest)

A modest effect on its own, but one trial found 500 mg of standardized extract twice daily equally effective as 12.5–25 mg of captopril twice daily, with an average reduction of 5–13 mmHg.[3][4][5]

🎯

ACE Inhibition

Secoiridoids — oleuropein, ligustroside, excelcioside, and related compounds — inhibit angiotensin-converting enzyme.[6][7]

⚖️

Beta-Blocker & Calcium-Channel Blocker Effect

A modest beta-blocking and calcium-channel-blocking effect, demonstrated in pharmacological screening.[8][9]

🫀

Coronary Vasodilator & Cardioprotective

Increases coronary blood flow; cardioprotective through combined antioxidant, antihypertensive, hypoglycemic, and hypocholesterolemic effects.[10]

🩸

Antiatherogenic

Documented antiatherogenic activity for olive leaf compounds.[11]

🔥

Myocardial Anti-Inflammatory Protection

Protects myocardial function via anti-inflammatory, antiatherosclerotic, anti-ischemic, and hypolipidemic effects.[12]

🧫

Cardiomyocyte Protection

Documented protective effect on heart muscle cells (cardiomyocytes).[13]

🩹

Vascular Function Protection

Protective effect on vascular function, documented separately from its cardiomyocyte and antiatherogenic effects.[14]

💓

Anti-Arrhythmic (Bradycardic)

Oleanolic acid produces an anti-arrhythmic, heart-rate-slowing effect via beta-adrenergic antagonism, with negative chronotropic and positive inotropic effects.[15]

🧬

Polyphenol Synergy (Cardioprotective)

An ethanolic leaf extract protects cardiomyocyte viability more effectively than any of its individual phenolic compounds alone.[13]

🩸

Antiplatelet Aggregation (Leaf)

Inhibits platelet aggregation.[16]

😌

Spasmolytic

Documented antispasmodic activity for the leaf.

⚗️

Antioxidant

Documented antioxidant activity across multiple independent studies of leaf phenolics.[17][18]

🩺

Hypocholesterolemic & Antidiabetic

Hypocholesterolemic, hypotriglyceridemic, and antidiabetic effects, especially pronounced in the wild oleaster variety; hypoglycemic activity confirmed in rats.[19][20]

🍯

Antidiabetic — Multiple Mechanisms

Alpha-amylase and alpha-glucosidase inhibition (luteolin); insulin-release potentiation and increased peripheral glucose uptake (oleuropeoside, 16 mg/kg); superior to glibenclamide in a rat model.[21][22][23][24]

📉

Improves Insulin Sensitivity

Improves insulin sensitivity and reduces metabolic syndrome risk in a randomized human trial of middle-aged overweight men.[25]

🦠

Antibacterial & Antifungal

Antibacterial and antifungal activity attributed to the leaf's phenolic derivatives.[26]

🫘

Nephroprotective

Documented nephroprotective activity for olive leaf.[27]

❓

Uncertain
Thyroid Stimulant

A pituitary-independent thyroid-stimulating activity has been proposed, but the primary source itself flags this finding with a question mark.[28]

🫒

Olive Oil
Anti-Inflammatory (Ibuprofen-Like)

Oleocanthal in olive oil produces anti-inflammatory activity comparable to ibuprofen.[29]

🩸

Olive Oil
Antiplatelet Aggregation

Phenolic components of olive oil inhibit platelet aggregation and eicosanoid production.[30]

🦴

Olive Oil
Anti-Osteoporotic (Animal)

Reduces ovariectomy-induced osteoporosis in a rat model.[31]

Clinical Applications

Clinical Indications

From mainstream phytotherapy uses to the more specialized gemmotherapy (bud extract) tradition.

🩺
Hypertension
Primary Indication
  • Via beta-blocker, calcium-channel-blocker, and ACE-inhibiting effects. [8][9][6]
  • EMA-recognized efficacy at 400 mg/day of dry extract. [32]
🩸
Circulatory & Metabolic
Phytotherapy
  • Venous circulatory disorders.
  • Mild diabetes (oleuropein, hydroxytyrosol) and reduction of diabetes-related complications (oleuropein). [33][34]
🌱
Gemmotherapy (Bud)
Standalone Use
  • Metabolic syndrome.
  • Non-thrombotic cerebral atherosclerosis, diabetic arteritis, gangrene. [35]
🤝
Gemmotherapy (Bud) — Combinations
With Other Plant Buds
  • With Prunus amygdalus (almond): phobic neurosis (agoraphobia, claustrophobia), obsessive-compulsive neurosis.
  • With Ficus carica (fig): facial neuralgia.
Pharmacodynamics

Mode of Action

The antihypertensive effect is attributed to three converging mechanisms.

⚖️

Beta-Blocker Effect

A modest beta-adrenergic blocking effect contributes to blood pressure reduction.[8][9]

🧪

Calcium-Channel Antagonism

Demonstrated calcium-channel-blocking activity, a second contributing mechanism to the antihypertensive effect.[8][9]

🎯

ACE Inhibition

Inhibition of angiotensin-converting enzyme by leaf secoiridoids forms the third mechanism.[6]

Clinical Safety

Safety & Precautions

One of the cleaner safety profiles among the plants in this series, with only mild theoretical interaction concerns.

⚠️

Adverse Effects

  • No adverse effects or interactions are documented in the primary phytotherapy literature for this plant.
  • Very mild pharmacokinetic interaction: oleuropein shows weak inhibition of the CYP1A2 liver enzyme. [36]
  • Weak CYP3A4 inhibition by maslinic acid, with low associated drug-interaction risk. [37]
🚫

Precautions

  • Potential pharmacodynamic interactions with antidiabetic and antihypertensive medications, given olive leaf's own effects on blood sugar and blood pressure.
  • Monitor blood pressure and blood glucose if combining with prescribed medication for either condition.
Clinical Disclaimer: This monograph is for educational and professional reference only. It does not constitute medical advice, diagnosis, or treatment guidance. Always consult a qualified healthcare provider before initiating any phytotherapeutic regimen, particularly if you are pregnant, breastfeeding, or taking antihypertensive or antidiabetic medication.
Common Questions

Frequently Asked Questions

What is olive leaf extract used for?
Olive leaf is studied primarily for mild hypertension, where a standardized extract has shown effects comparable to low-dose captopril, as well as for mild diabetes, venous circulatory disorders, and broader cardioprotective and antioxidant support.
What is the recommended dose of olive leaf extract?
The standard phytotherapy dose is 400 to 500 mg of dry extract taken twice daily (800 mg to 1 g per day), in capsule form. A leaf decoction (30 g per liter, one liter per day) is also traditionally used. Always confirm dosing with a healthcare provider.
What is oleuropein?
Oleuropein is the principal secoiridoid compound in olive leaf, making up a significant portion of the leaf's dry weight. It is considered the main compound behind olive leaf's antihypertensive, antioxidant, and antidiabetic effects, and most standardized extracts are measured by their oleuropein content.
Can olive leaf extract lower blood pressure as well as medication?
One randomized trial found 500 mg of standardized olive leaf extract twice daily comparably effective to 12.5 to 25 mg of captopril twice daily in patients with stage-1 hypertension, with an average reduction of 5 to 13 mmHg. This is a single trial rather than a large, definitive evidence base, so it should be discussed with a healthcare provider rather than used as a substitute for prescribed medication.
Does olive leaf extract interact with medications?
No significant adverse effects or interactions are documented in the primary literature reviewed, and laboratory studies show only very mild effects on liver enzymes (weak CYP1A2 and CYP3A4 inhibition), suggesting low interaction risk. That said, because olive leaf has its own blood-sugar-lowering and blood-pressure-lowering effects, a theoretical additive effect with diabetes and hypertension medications is reasonable to discuss with a healthcare provider.
Is olive leaf the same as olive oil?
No. Olive leaf extract is made from the dried leaves of the olive tree and is rich in oleuropein, a compound barely present in the fruit. Olive oil is pressed from the fruit itself and contains a different profile of compounds, including oleocanthal, though both are credited with anti-inflammatory and antiplatelet effects.
Can olive leaf help with diabetes?
Animal and laboratory research points to several antidiabetic mechanisms, including alpha-glucosidase inhibition, enhanced glucose-induced insulin release, and improved insulin sensitivity in a human trial of middle-aged overweight men. The primary source describes this as appropriate for mild diabetes rather than a replacement for standard diabetes treatment.
Primary Literature

Bibliography

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↑ 2. Stiti N, Hartmann MA. Nonsterol Triterpenoids as Major Constituents of Olea europaea. J Lipids. 2012;2012:476595. PubMed PMID:22523691 →
↑ 3. Khayyal MT, el-Ghazaly MA, Abdallah DM, Nassar NN, Okpanyi SN, Kreuter MH. Blood pressure lowering effect of an olive leaf extract (Olea europaea) in L-NAME induced hypertension in rats. Arzneimittelforschung. 2002;52(11):797-802. PubMed PMID:12489249 →
↑ 4. Susalit E, Agus N, Effendi I, Tjandrawinata RR, Nofiarny D, Perrinjaquet-Moccetti T, Verbruggen M. Olive (Olea europaea) leaf extract effective in patients with stage-1 hypertension: comparison with Captopril. Phytomedicine. 2011 Feb 15;18(4):251-258. PubMed PMID:21036583 →
↑ 5. Perrinjaquet-Moccetti T, Busjahn A, Schmidlin C, Schmidt A, Bradl B, Aydogan C. Food supplementation with an olive (Olea europaea L.) leaf extract reduces blood pressure in borderline hypertensive monozygotic twins. Phytother Res. 2008 Sep;22(9):1239-1242. PubMed PMID:18729245 →
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↑ 7. Lockyer S, Rowland I, Spencer JPE, Yaqoob P, Stonehouse W. Impact of phenolic-rich olive leaf extract on blood pressure, plasma lipids and inflammatory markers: a randomised controlled trial. Eur J Nutr. 2017 Jun;56(4):1421-1432. PubMed PMID:26951205 →
↑ 8. Scheffler A, Rauwald HW, Kampa B, Mann U, Mohr FW, Dhein S. Olea europaea leaf extract exerts L-type Ca(2+) channel antagonistic effects. J Ethnopharmacol. 2008 Nov 20;120(2):233-240. PubMed PMID:18790040 →
↑ 9. Rauwald HW, Brehm O, Odenthal KP. Screening of nine vasoactive medicinal plants for their possible calcium antagonistic activity. Strategy of selection and isolation for the active principles of Olea europaea and Peucedanum ostruthium. Phytother Res. 1994 May;8(3):135-140.
↑ 10. Vogel P, Kasper Machado I, Garavaglia J, Zani VT, de Souza D, Morelo Dal Bosco S. Polyphenols benefits of olive leaf (Olea europaea L) to human health. Nutr Hosp. 2014 Dec 17;31(3):1427-1433. PubMed PMID:25726243 →
↑ 11. El SN, Karakaya S. Olive tree (Olea europaea) leaves: potential beneficial effects on human health. Nutr Rev. 2009 Nov;67(11):632-638. PubMed PMID:19906250 →
↑ 12. Efentakis P, Iliodromitis EK, Mikros E, Papachristodoulou A, Dagres N, Skaltsounis AL, Andreadou I. Effects of the olive tree leaf constituents on myocardial oxidative damage and atherosclerosis. Planta Med. 2015 Jun;81(8):648-654. PubMed PMID:26018920 →
↑ 13. Bali EB, Ergin V, Rackova L, Bayraktar O, Küçükboyaci N, Karasu Ç. Olive leaf extracts protect cardiomyocytes against 4-hydroxynonenal-induced toxicity in vitro: comparison with oleuropein, hydroxytyrosol, and quercetin. Planta Med. 2014 Aug;80(12):984-992. PubMed PMID:25098929 →
↑ 14. Lockyer S, Corona G, Yaqoob P, Spencer JP, Rowland I. Secoiridoids delivered as olive leaf extract induce acute improvements in human vascular function and reduction of an inflammatory cytokine: a randomised, double-blind, placebo-controlled, cross-over trial. Br J Nutr. 2015 Jul 14;114(1):75-83. PubMed PMID:26051429 →
↑ 15. Javidanpour S, Dianat M, Aliakbari FR, Sarkaki A. The effects of olive leaf extract and 28 days forced treadmill exercise on electrocardiographic parameters in rats. J Res Med Sci. 2018;23:108. DOI:10.4103/jrms.JRMS_517_18 →
↑ 16. Singh I, Mok M, Christensen AM, Turner AH, Hawley JA. The effects of polyphenols in olive leaves on platelet function. Nutr Metab Cardiovasc Dis. 2008 Feb;18(2):127-132. PubMed PMID:17346951 →
↑ 17. Benavente-García O, Castillo J, Lorente J, Ortuño A, Del Rio JA. Antioxidant activity of phenolics extracted from Olea europaea L. leaves. Food Chemistry. 2000 Mar;68(4):457-462.
↑ 18. Lkrik A, Souidi K, Martin P. Effet des polyphénols extraits à partir des tourteaux et feuilles de l'olivier (Olea europaea L) sur la stabilité oxydative de l'huile d'olive. Full text →
↑ 19. Bennani-Kabchi N, Fdhil H, Cherrah Y, Kehel L, el Bouayadi F, Amarti A, Saïdi M, Marquié G. Effects of Olea europea var. oleaster leaves in hypercholesterolemic insulin-resistant sand rats. Therapie. 1999 Nov-Dec;54(6):717-723. PubMed PMID:10709446 →
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↑ 21. Eddouks M, Chattopadhyay D. Phytotherapy in the Management of Diabetes and Hypertension. Bentham Science Publishers, 2012.
↑ 22. Gonzalez M, Zarzuelo A, Gamez MJ, Utrilla MP, Jimenez J, Osuna I. Hypoglycemic activity of olive leaf. Planta Med. 1992 Dec;58(6):513-515. PubMed PMID:1484890 →
↑ 23. Eidi A, Eidi M, Darzi R. Antidiabetic effect of Olea europaea L. in normal and diabetic rats. Phytother Res. 2009 Mar;23(3):347-350. PubMed PMID:18844257 →
↑ 24. Sato H, Genet C, Strehle A, Thomas C, Lobstein A, Wagner A, Mioskowski C, Auwerx J, Saladin R. Anti-hyperglycemic activity of a TGR5 agonist isolated from Olea europaea. Biochem Biophys Res Commun. 2007 Nov 3;362(4):793-798. PubMed PMID:17825251 →
↑ 25. de Bock M, Derraik JG, Brennan CM, Biggs JB, Morgan PE, Hodgkinson SC, Hofman PL, Cutfield WS. Olive (Olea europaea L.) leaf polyphenols improve insulin sensitivity in middle-aged overweight men: a randomized, placebo-controlled, crossover trial. PLoS One. 2013;8(3):e57622. PubMed PMID:23516412 →
↑ 26. Pereira AP, Ferreira IC, Marcelino F, Valentão P, Andrade PB, Seabra R, Estevinho L, Bento A, Pereira JA. Phenolic compounds and antimicrobial activity of olive (Olea europaea L. Cv. Cobrançosa) leaves. Molecules. 2007 May 26;12(5):1153-1162. PubMed PMID:17873849 →
↑ 27. Azab AE, Albasha MO, Elsayed ASI. Prevention of Nephropathy by Some Natural Sources of Antioxidants. Yangtze Medicine. 2017;1:235-266. DOI:10.4236/ym.2017.14023 →
↑ 28. Al-Qarawi AA, Al-Damegh MA, ElMougy SA. Effect of freeze dried extract of Olea europaea on the pituitary-thyroid axis in rats. Phytother Res. 2002 May;16(3):286-287. PubMed PMID:12164280 →
↑ 29. Beauchamp GK, Keast RS, Morel D, Lin J, Pika J, Han Q, Lee CH, Smith AB, Breslin PA. Phytochemistry: ibuprofen-like activity in extra-virgin olive oil. Nature. 2005 Sep 1;437(7055):45-46. PubMed PMID:16136122 →
↑ 30. Petroni A, Blasevich M, Salami M, Papini N, Montedoro GF, Galli C. Inhibition of platelet aggregation and eicosanoid production by phenolic components of olive oil. Thromb Res. 1995 Apr 15;78(2):151-160. PubMed PMID:7482432 →
↑ 31. Saleh NK, Saleh HA. Olive Oil effectively mitigates Ovariectomy induced Osteoporosis In Rats. BMC Complement Altern Med. 2011 Feb 4;11:10. Full text →
↑ 32. Committee on Herbal Medicinal Products (HMPC), European Medicines Agency. Assessment report on Olea europaea L., folium. EMA/HMPC/359236/2016, 31 January 2017. EMA Assessment Report →
↑ 33. Jemai H, El Feki A, Sayadi S. Antidiabetic and antioxidant effects of hydroxytyrosol and oleuropein from olive leaves in alloxan-diabetic rats. J Agric Food Chem. 2009 Oct 14;57(19):8798-8804. PubMed PMID:19725535 →
↑ 34. Al-Azzawie HF, Alhamdani MS. Hypoglycemic and antioxidant effect of oleuropein in alloxan-diabetic rabbits. Life Sci. 2006 Feb 16;78(12):1371-1377. PubMed PMID:16236331 →
↑ 35. Pol H. Gemmothérapie, thérapeutique par les extraits embryonnaires végétaux. Self-published, Brussels, 1982.
↑ 36. Stupans I, Murray M, Kirlich A, Tuck KL, Hayball PJ. Inactivation of cytochrome P450 by the food-derived complex phenol oleuropein. Food Chem Toxicol. 2001;39(11):1119-1124. DOI:10.1016/S0278-6915(01)00060-6 →
↑ 37. Sun M, Tang Y, Ding T, Liu M, Wang X. Investigation of cytochrome P450 inhibitory properties of maslinic acid, a bioactive compound from Olea europaea L., and its structure–activity relationship. Phytomedicine. 2015;22(1):56-65. DOI:10.1016/j.phymed.2014.10.003 →

Additional Reference Literature

Bruneton J. Pharmacognosie, Phytochimie, Plantes médicinales. Ed. Tec & Doc, 1997. p. 487.
Rovellini P, Cortesi N, Fedeli E. Analysis of flavonoids from Olea Europaea by HPLC-UV and HPLC-electrospray-MS. Rivista Italiana delle Sostanze Grasse. 1997;74(7):273-279.
Ficarra P, Ficarra R, de Pasquale A, Monforte MT, Calabrò ML. HPLC analysis of oleuropein and some flavonoids in leaf and bud of Olea europaea L. Farmaco. 1991 Jun;46(6):803-815. PubMed PMID:1772565 →
Reyes FJ, Centelles JJ, Lupiáñez JA, Cascante M. (2α,3β)-2,3-Dihydroxyolean-12-en-28-oic acid, a new natural triterpene from Olea europea, induces caspase dependent apoptosis selectively in colon adenocarcinoma cells. FEBS Lett. 2006 Nov 27;580(27):6302-6310.