ÖkeAltuntaş F, Demi Rtaş I (2017) Real-time cell analysis of the cytotoxicity of Origanum
acutidens essential oil on HT-29 and HeLa cell lines. Turk J Pharm Sci 14:29–33
Ozkan A, Erdogan A (2012) A comparative study of the antioxidant/prooxidant effects of
carvacrol and thymol at various concentrations on membrane and DNA of parental and drug
resistant H1299 cells. Nat Prod Commun 7:1557–1560
Pakdemi rli A, Karaca C, Sever T, Daskin E, Leblebi Ci A, Yi Ği Tbaşi T, Başbinar Y (2020)
Carvacrol alters soluble factors in HCT-116 and HT-29 cell lines. Turk. J. Med. Sci. 50:271–
276
Patil JU, Suryawanshi KC, Patil PB, Chaudhary SR, Pawar NS (2010) Synthesis and antibacterial
activity of carvacryl ethers. J Asian Nat Prod Res 12:129–133
Pilau MR, Alves SH, Weiblen R, Arenhart S, Cueto AP, Lovato LT (2011) Antiviral activity of
the Lippia graveolens (Mexican oregano) essential oil and its main compound carvacrol
against human and animal viruses. Braz J Microbiol. 42(4):1616–1624
Potočnjak I, Gobin I, Domitrović R (2018) Carvacrol induces cytotoxicity in human cervical
cancer cells but causes cisplatin resistance: Involvement of MEK-ERK activation. Phytother.
Res. 32:1090–1097
Ramak P, Kazempour Osaloo S, Ebrahimzadeh H, Sharifi M, Behmanesh M (2013) Inhibition of
the mevalonate pathway enhances carvacrol biosynthesis and DXR gene expression in shoot
cultures of Satureja khuzistanica Jamzad. J Plant Physiol 170:1187–1193
Rattanachaikunsopon P, Phumkhachorn P (2010) Assessment of factors influencing antimicrobial
activity of carvacrol and cymene against Vibrio cholerae in food. J Biosci Bioeng 110:614–619
Ravishankar S, Zhu L, Reyna-Granados J, Law B, Joens L, Friedman M (2010) Carvacrol and
cinnamaldehyde inactivate antibiotic-resistant Salmonella enterica in buffer and on celery and
oysters. J Food Prot 73:234–240
Report Final (2016) The Safety Assessment of Sodium p-chloro- m-cresol, p-chloro- m-cresol,
chlorothymol, mixed cresols, m-Cresol, o-cresol, p-cresol, isopropyl cresols, thymol, ocymen-5-ol, and carvacrol. Int. J. Toxicol. 25:29–127
Rohmer M, Knani M, Simonin P, Sutter B, Sahm H (1993) Isoprenoid biosynthesis in bacteria: a
novel pathway for the early steps leading to isopentenyldiphosphate. Biochem J 295:517–524
Samarghandian S, Farkhondeh T, Samini F, Borji A (2016) Protective effects of carvacrol against
oxidative stress induced by chronic stress in rat’s brain, liver, and kidney. Biochem Res Int.
2016:1–7
Sharifi-Rad M, Varoni EM, Iriti M, Martorell M, Setzer WN, del Mar Contreras M, Salehi B,
Soltani-Nejad A, Rajabi S, Tajbakhsh M, Sharifi-Rad J (2018) Carvacrol and human health: A
comprehensive review, Phyto Res. 1–13
Shimoda K, Kondo Y, Nishida T, Hamada H, Nakajima N, Hamada H (2006) Biotransformation
of thymol, carvacrol, and eugenol by cultured cells of Eucalyptus perriniana. Phytochemistry
67:2256–2261
Sim JXF, Khazandi M, Chan WY, Trott DJ, Deo P (2019) Antimicrobial activity of thyme oil,
oregano oil, thymol and carvacrol against sensitive and resistant microbial isolates from dogs
with Otitis externa. Vet Dermatol 30:524
Slamenova D, Kozics K, Hunakova L, Melusova M, Navarova J, Horvathova E (2013)
Comparison of biological processes induced in HepG2 cells by tert-butyl hydroperoxide
(t-BHP) and hydroperoxide(H 2 O 2 ): The influence of carvacrol. Mutat. Res. Genet. Toxicol.
Environ. 757:15–22
Takahashi S, Kuzuyama T, Watanabe H, Seto H (1998) A 1-deoxy-D-xylulose 5-phosphate
reductoisomerase catalyzing the formation of 2-C-methyl-D-erythritol 4-phosphate in an
alternative nonmevalonate pathway for terpenoid biosynthesis. Proc Natl Acad Sci USA
95:9879–9884
Teissedre PL, Waterhouse AL (2008) Inhibition of oxidation of human low-density lipoproteins by
phenolic substances in different essential oils varieties. J Agric Food Chem 48:3801–3805
13 Carvacrol (Origanum vulgare): Therapeutic Properties …
461
acutidens essential oil on HT-29 and HeLa cell lines. Turk J Pharm Sci 14:29–33
Ozkan A, Erdogan A (2012) A comparative study of the antioxidant/prooxidant effects of
carvacrol and thymol at various concentrations on membrane and DNA of parental and drug
resistant H1299 cells. Nat Prod Commun 7:1557–1560
Pakdemi rli A, Karaca C, Sever T, Daskin E, Leblebi Ci A, Yi Ği Tbaşi T, Başbinar Y (2020)
Carvacrol alters soluble factors in HCT-116 and HT-29 cell lines. Turk. J. Med. Sci. 50:271–
276
Patil JU, Suryawanshi KC, Patil PB, Chaudhary SR, Pawar NS (2010) Synthesis and antibacterial
activity of carvacryl ethers. J Asian Nat Prod Res 12:129–133
Pilau MR, Alves SH, Weiblen R, Arenhart S, Cueto AP, Lovato LT (2011) Antiviral activity of
the Lippia graveolens (Mexican oregano) essential oil and its main compound carvacrol
against human and animal viruses. Braz J Microbiol. 42(4):1616–1624
Potočnjak I, Gobin I, Domitrović R (2018) Carvacrol induces cytotoxicity in human cervical
cancer cells but causes cisplatin resistance: Involvement of MEK-ERK activation. Phytother.
Res. 32:1090–1097
Ramak P, Kazempour Osaloo S, Ebrahimzadeh H, Sharifi M, Behmanesh M (2013) Inhibition of
the mevalonate pathway enhances carvacrol biosynthesis and DXR gene expression in shoot
cultures of Satureja khuzistanica Jamzad. J Plant Physiol 170:1187–1193
Rattanachaikunsopon P, Phumkhachorn P (2010) Assessment of factors influencing antimicrobial
activity of carvacrol and cymene against Vibrio cholerae in food. J Biosci Bioeng 110:614–619
Ravishankar S, Zhu L, Reyna-Granados J, Law B, Joens L, Friedman M (2010) Carvacrol and
cinnamaldehyde inactivate antibiotic-resistant Salmonella enterica in buffer and on celery and
oysters. J Food Prot 73:234–240
Report Final (2016) The Safety Assessment of Sodium p-chloro- m-cresol, p-chloro- m-cresol,
chlorothymol, mixed cresols, m-Cresol, o-cresol, p-cresol, isopropyl cresols, thymol, ocymen-5-ol, and carvacrol. Int. J. Toxicol. 25:29–127
Rohmer M, Knani M, Simonin P, Sutter B, Sahm H (1993) Isoprenoid biosynthesis in bacteria: a
novel pathway for the early steps leading to isopentenyldiphosphate. Biochem J 295:517–524
Samarghandian S, Farkhondeh T, Samini F, Borji A (2016) Protective effects of carvacrol against
oxidative stress induced by chronic stress in rat’s brain, liver, and kidney. Biochem Res Int.
2016:1–7
Sharifi-Rad M, Varoni EM, Iriti M, Martorell M, Setzer WN, del Mar Contreras M, Salehi B,
Soltani-Nejad A, Rajabi S, Tajbakhsh M, Sharifi-Rad J (2018) Carvacrol and human health: A
comprehensive review, Phyto Res. 1–13
Shimoda K, Kondo Y, Nishida T, Hamada H, Nakajima N, Hamada H (2006) Biotransformation
of thymol, carvacrol, and eugenol by cultured cells of Eucalyptus perriniana. Phytochemistry
67:2256–2261
Sim JXF, Khazandi M, Chan WY, Trott DJ, Deo P (2019) Antimicrobial activity of thyme oil,
oregano oil, thymol and carvacrol against sensitive and resistant microbial isolates from dogs
with Otitis externa. Vet Dermatol 30:524
Slamenova D, Kozics K, Hunakova L, Melusova M, Navarova J, Horvathova E (2013)
Comparison of biological processes induced in HepG2 cells by tert-butyl hydroperoxide
(t-BHP) and hydroperoxide(H 2 O 2 ): The influence of carvacrol. Mutat. Res. Genet. Toxicol.
Environ. 757:15–22
Takahashi S, Kuzuyama T, Watanabe H, Seto H (1998) A 1-deoxy-D-xylulose 5-phosphate
reductoisomerase catalyzing the formation of 2-C-methyl-D-erythritol 4-phosphate in an
alternative nonmevalonate pathway for terpenoid biosynthesis. Proc Natl Acad Sci USA
95:9879–9884
Teissedre PL, Waterhouse AL (2008) Inhibition of oxidation of human low-density lipoproteins by
phenolic substances in different essential oils varieties. J Agric Food Chem 48:3801–3805
13 Carvacrol (Origanum vulgare): Therapeutic Properties …
461