A pharmacological and phytochemical overview on Satureja

Abstract

Context: Satureja (Lamiaceae) species are used as flavoring compounds in food, pharmaceutical, and cosmetic industries because of its sweetness and simple cultivation characteristics. They have traditionally been used as muscle pain relievers, tonic, and carminative agents to treat stomach and intestinal disorders such as cramps, nausea, indigestion, and diarrhea, due to their considerable phytochemical characteristics.

Objective: This review evaluates some information published since 1989 on Satureja genus from a systematic perspective in terms of its pharmacological and phytochemical characteristics.

Materials and methods: Web of Science, PubMed, Scopus, and Google Scholar databases were searched up to March 2014. “Satureja” was used as a research term without narrowing or limiting research elements. After obtaining all reports from database (a total number is about 637), the papers were carefully analyzed in order to find data related to the topic of this review.

Results: In this review, 453 reports were used which were published between 1989 and 2014. The study was compiled into two sections (Pharmacology and Phytochemistry). In the pharmacology section, more than 50 different activities were studied. In the second section, in addition to other compounds and inorganic substances, volatiles, phenolic acids, and flavonoids were discussed.

Discussion and conclusion: According to our contemporary information, Satureja species have been evaluated for their wide range of biological activities. However, a small part of these studies have been carried out on the active principles. Therefore, in the future, more studies should be carried out to identify responsible phytochemicals for the various activities.

• Biological activity
• essential oil
• extract
• phytochemistry

Introduction

Selection of medicinal plants requires experience. Statistical studies that were carried out in Australia and US indicated that almost 48.5% and 34% of individuals had used at least one unconventional therapy, respectively (Kim et al., 2007; Momtaz & Abdollahi, 2010). In developing countries, about 65–80% of the world population depends primarily on plants for their health care due to poverty and lack of access to modern medicine (Calixto, 2005; Momtaz & Abdollahi, 2010).

It is well known that in many developing countries, people are suffering from lack of safe modern drugs. Therefore, evaluation of effective plants for the treatment of diseases such as diabetes has been recommended (Momtaz & Abdollahi, 2010). In the last decade, many researchers have reported efficacy of herbal medicines in many diseases. According to statistical data reported several times (Kim et al., 2007; Momtaz & Abdollahi, 2010), in currently prescribed synthetic drugs, approximately 25% of active compounds were first identified in plant resources. Moreover, 20 000 plants have been used for medicinal purposes some of which around 4000 have been used commonly. Among them, 10% have commercial importance currently (Kim et al., 2007; Momtaz & Abdollahi, 2010).

The genus Satureja (Lamiaceae) was first named by Roman writer Pliny. The name comes from Latin “satureia”. It means “herb of satyrs”; because of that reason its cultivation was banned in monasteries (Zavatti et al., 2011). This genus contains about 200 species of aromatic herbs and shrubs which grown mainly in the Middle East, Mediterranean region to Europe, West Asia, North Africa, the Canary Islands, and South America. More than 30 species of this genus are distributed in eastern parts of Mediterranean area (Cantino et al., 1992; Momtaz & Abdollahi, 2010). Due to its sweetness and simple cultivation characteristics, they are used as flavoring compounds in food, pharmaceutical, and cosmetic industries. They have traditionally been used as muscle pain reliever, tonic, and carminative agents in order to treat stomach and intestinal disorders such as cramps, nausea, indigestion, and diarrhea (Momtaz & Abdollahi, 2010; Zargari, 1990).

Pharmacology

Antimicrobial activity

Antimicrobial activity potential of the members of Satureja genus is presented in Table 1. As can be seen in the table, 191 studies concerning the antimicrobial activity of 37 different plant species have been reported elsewhere.

Table 1. Biological activities of Satureja species.

Medicinal use	Plant species	Reference
Antimicrobial activity	S. abyssinica subsp. abyssinica	Tolossa et al. (2007)
	S. aintabensis	Askun et al. (2013) and Azaz et al. (2005)
	S. bachtiarica	Ahanjan et al. (2011a, 2011b), Goudarzi et al. (2011), Hadian et al. (2012), and Pirbalouti et al. (2010, 2011a, 2011b)
	S. biflora	Matasyoh et al. (2007) and Vagionas et al. (2007)
	S. boissieri	Azaz et al. (2002)
	S. brownei	Caceres et al. (1991)
	S. cilicica	Askun et al. (2012) and, Kiran et al. (2010)
	S. coerulea	Askun et al. (2012) and Azaz et al. (2002)
	S. cuneifolia	Azaz et al. (2005), Baydar et al. (2004), Biavati et al. (2004), Kan et al. (2006), Kosar et al. (2008), Menkovic et al. (2007), Mirjana et al. (2004), Oke et al. (2009a, 2009b), Orhan et al. (2011, 2012), Savikin et al. (2010), Skocibusic and Bezic (2004), Stefanini et al. (2006), and Tommasi et al. (2009)
	S. edmondi	Masumeh (2006)
	S. hortensis	Abdollahi et al. (2010, 2011, 2012), Adiguzel et al. (2007), Aksu and Ozer (2013), Alizadeh-Salteh et al. (2010b), Anghel et al. (2013), Askun et al. (2008), Azaz et al. (2005), Azizi et al. (2008), Bouari et al. (2011), Boyraz and Ozcan (2006), Deans and Svoboda (1989), Dikbas et al. (2008, 2010, 2012), Djenane et al. (2011a), Fit et al. (2009), Gulluce et al. (2003), Gumus (2010), Gursoy et al. (2009), Hosseini et al. (2013), Karami-Osboo et al. (2010), Kiran et al. (2010), Kizil and Uyar (2006), Kotan et al. (2012, 2013), Mahboubi and Kazempour (2009), Mihajilov-Krstev et al. (2009a, 2009b, 2010), Nabigol (2011), Nedorostova et al. (2011), Oussalah et al. (2006b), Ozcan et al. (2008), Razzaghi-Abyaneh et al. (2008), Rezvanpanah et al. (2011), Sagdic et al. (2010, 2013), Saharkhiz et al. (2011), Sahin et al. (2003), Shirzad et al. (2011), Shojaee-Aliabadi et al. (2013), Stompor-Chrzan (2003), Tuchila et al. (2008), Venskutonis et al. (2010), and Yazici et al. (2011)
	S. horvatii	Lakusic et al. (2008)
	S. icarica	Askun et al. (2012), Azaz et al. (2002), and Dulger and Hacioglu (2008)
	S. inontana	Oussalah et al. (2006a)
	S. intermedia	Sadeghi et al. (2013)
	S. khuzestanica	Amanlou et al. (2004), Hadian et al. (2012), Nabigol (2011), Pirbalouti and Moalem (2013), Pirbalouti et al. (2011b), Rezvanfar et al. (2008a), Sadeghi-Nejad et al. (2010), Saei-Dehkordi et al. (2012), Taban et al. (2013), Yousefzadi et al. (2014), and Zarrin et al. (2010)
	S. kitaibelii	Kundakovic et al. (2011), Mihajilov-Krstev et al. (2011), Zivkovic et al. (2009)
	S. macrantha	Azaz et al. (2005)
	S. masukensis	Vagionas et al. (2007)
	S. montana	Bezbradica et al. (2005), Carraminana et al. (2008), Cavar et al. (2008), Cetojevic-Simin et al. (2008), Chao et al. (2000), Christian and Goggi (2008), Ciani et al. (2000), Damjanovic-Vratnica et al. (2011), de Oliveira et al. (2011), Djenane et al. (2011b, 2012), Fraternale et al. (2007), Giordani et al. (2004), Haloci et al. (2011), Lopez-Reyes et al. (2010), Marin et al. (2012), Miladi et al. (2013), Mirjana and Nada (2004), Mohacsi-Farkas et al. (2003), Nedorostova et al. (2009, 2011), Oussalah et al. (2006b, 2007), Oussalaili et al. (2007), Panizzi et al. (1993), Penalver et al. (2005), Ratajac et al. (2011), Rota et al. (2004), Serrano et al. (2011), Sfeir et al. (2013), Silva et al. (2009), Skocibusic and Bezic (2004), Stefanini et al. (2006), Tampieri et al. (2005), and Turgis et al. (2012)
	S. montana subsp. kitaibelii	Cetkovic et al. (2007a)
	S. mutica	Hadian et al. (2012) and Taghvaei et al. (2009)
	S. odora	Fuselli et al. (2008)
	S. parnassica	Chorianopoulos et al. (2006a)
	S. parnassica subsp. parnassica	Chorianopoulos et al. (2004) and Tzakou and Skaltsa (2003)
	S. parvifolia	Feresin et al. (2001), Hernandez et al. (2000), Lima et al. (2011), Luna et al. (2008), Zygadlo and Grosso (1995)
	S. pilosa	Azaz et al. (2002)
	S. pomifera subsp. calycina	Glamoclija et al. (2006)
	S. pseudosimensis	Tchoumbougnang et al. (2009) and Vagionas et al. (2007)
	S. punctata	Tchoumbougnang et al. (2009)
	S. robusta	Tchoumbougnang et al. (2009)
	S. sahendica	Taherpour et al. (2008) and Yousefzadi et al. (2012)
	S. spicigera	Eftekhar et al. (2009) and Kotan et al. (2010), Nabigol (2011), Sariboga and Sariboga (2011)
	S. spinosa	Chorianopoulos et al. (2004, 2006b)
	S. subspicata	Cavar et al. (2008) and Skocibusic et al. (2006)
	S. thymbra	Azaz et al. (2005), Boziaris et al. (2011), Chorianopoulos et al. (2004, 2006a, 2008), Giweli et al. (2012), Glamoclija et al. (2006), Goren et al. (2004), Gumus et al. (2010), Lebert et al. (2007), Markovic et al. (2011), Mullerriebau et al. (1995), Okmen et al. (2012), Piras et al. (2011), Sarac and Ugur (2008), Shimoni et al. (1993), Sokovic et al. (2002), Ugur et al. (2009), Vokou et al. (1993), and Yegen et al. (1992)
	S. wiedemanniana	Altundag and Aslim (2011), Baser et al. (2001), Yucel and Aslim (2011), and Yucel et al. (2009)
	Antioxidant activity	S. abyssinica subsp. abyssinica	Tolossa et al. (2007)
S. bachtiarica		Hashemi et al. (2011)
S. cilicica		Ozkan et al. (2007)
S. cuneifolia		Dorman et al. (2004), Eminagaoglu et al. (2007), Kosar et al. (2003, 2008), Kulisic et al. (2005), Oke et al. (2009a, 2009b), and Stefanini et al. (2006)
S. delpozoi		Jordan et al. (2010)
S. forbesii		Ortet et al. (2009)
S. laevigata		Villa-Ruano et al. (2013)
S. hortensis		Alinkina et al. (2013), Alizadeh (2011), Alizadeh et al. (2010a), Bandoniene et al. (2002), Deighton et al. (1994), Dikbas et al. (2011), Dorman and Hiltunen (2004), Esquivel et al. (1999), Exarchou et al. (2002), Gulluce et al. (2003), Hosseini et al. (2013), Inotai et al. (2012), Kalantzakis and Blekas (2006), Kosar et al. (2005), Madsen et al. (1996, 1998), Plander et al. (2012), Shojaee-Aliabadi et al. (2013), Trojakova et al. (2001), and Yesiloglu et al. (2013)
S. intricata		Jordan et al. (2010)
S. khuzestanica		Hashemi et al. (2012), Rezvanfar et al. (2008b, 2009, 2010), Saei-Dehkordi et al. (2012), Safarnavadeh and Rastegarpanah (2011), and Vosough-Ghanbari et al. (2010)
S. laxiflora		Sonboli et al. (2004)
S. macrosiphonia		Amiri (2011)
S. macrostema		Gutierrez (2013) and Gutierrez and Navarro (2010)
S. montana		Cavar et al. (2008), Chrpova et al. (2010), de Oliveira et al. (2012), Giao et al. (2009, 2010, 2012, 2013), Grosso et al. (2009b), Kulisic et al. (2005), Marin et al. (2012), Mercado-Mercado et al. (2013), Miladi et al. (2013), Perez-Cruz et al. (2013), Pestana et al. (2007), Prieto et al. (2007), Radonic and Milos (2003b), Serrano et al. (2011), Stefanini et al. (2006), Stoilova et al. (2008), and Vidic et al. (2010)
S. montana subsp. kitaibelii		Cetkovic et al. (2007a)
S. montana subsp. montana		Radonic and Milos (2003a)
S. mutica		Motamed and Naghibi (2010)
S. obovata		Jordan et al. (2010)
S. parvifolia		Cabana et al. (2013) and Dade et al. (2009), Zygadlo et al. (1995)
S. spicigera		Eminagaoglu et al. (2007)
S. subspicata		Cavar et al. (2008)
S. thymbra		Giweli et al. (2012), Lagouri and Boskou (1996), and Lagouri et al. (1993)
S. visianii		Vidic et al. (2010)
Cytotoxic activity		S. atropatana	Moradi et al. (2008)
	S. hortensis	Manjili et al. (2012)
	S. intermedia	Sadeghi et al. (2013)
	S. kitaibelii	Stanojkovic et al. (2013)
	S. khuzestanica	Yousefzadi et al. (2014)
	S. montana	Miladi et al. (2013)
	S. odora	Mongelli et al. (1996)
	S. parvifolia	Mongelli et al. (1996)
	S. punctata subsp. punctata	Tariku et al. (2010)
	S. sahendica	Yousefzadi et al. (2012)
	S. spicigera	Gohari et al. (2012)
	S. thymbra	Saab et al. (2012)
	Insecticidal activity / Fumigant  toxicity	S. hortensis	Aslan et al. (2004), Ebadollahi et al. (2012), Kim et al. (2012), Maedeh et al. (2011), Pavela (2008), Pavela et al. (2008, 2009), Sajfrtova et al. (2013), Tozlu et al. (2011), Yildirim et al. (2011), and Zandi-Sohani (2011)
S. montana		Faria et al. (2013) and Michaelakis et al. (2007)
S. parnassica subsp. parnassica		Michaelakis et al. (2007)
S. spicigera		Yildirim et al. (2011)
S. spinosa		Michaelakis et al. (2007)
S. thymbra		Ayvaz et al. (2009), Dell'Agli et al. (2012), Karaborklu et al. (2011), Karamaouna et al. (2013), Karpouhtsis et al. (1998), Michaelakis et al. (2007), and Sarac and Tunc (1995a, 1995b)
Acetylcholinesterase inhibitory  activity		S. cuneifolia	Orhan et al. (2008)
	S. khuzestanica	Basiri et al. (2007)
	S. montana	Silva et al. (2009)
	S. parvifolia	Cabana et al. (2013)
	S. thymbra	Ozturk (2012)
	Insect repellant activity	S. hortensis	Nottingham and Hardie (1993) and Nottingham et al. (1991)
S. khuzestanica		Kayedi et al. (2011, 2014)
S. montana		Picard et al. (2012)
S. parvifolia		Lima et al. (2011)
S. thymbra		Sarac and Tunc (1995a)
Antiviral activity		S. boliviana	Abad et al. (1999)
	S. cuneifolia	Orhan et al. (2012)
	S. montana	Yamasaki et al. (1998)
	S. montana subsp. variegata	Dunkic et al. (2010)
	S. thymbra	Loizzo et al. (2008b), Saab et al. (2012)
	Antinociceptive / Analgesic  activity	S. cuneifolia	Aydin et al. (1996)
S. hortensis		Hajhashemi et al. (2002, 2012)
S. khuzestanica		Amanlou et al. (2005)
S. thymbra		Karabay-Yavasoglu et al. (2006)
S. viminea		Suarez et al. (2003)
Antileishmanial activity		S. bachtiarica	Mohammadpour et al. (2012)
	S. hortensis	Manjili et al. (2012)
	S. khuzestanica	Kheirandish et al. (2011) and Sadeghi-Nejad et al. (2011)
	S. punctata subsp. punctata	Tariku et al. (2010)
	Genotoxic activity	S. hortensis	Panahi et al. (2009) and Zani et al. (1991)
S. montana		Zani et al. (1991)
S. thymbra		Karpouhtsis et al. (1998)
Anti-inflammatory activity		S. hortensis	Hajhashemi et al. (2002, 2012)
	S. khuzestanica	Amanlou et al. (2005) and Ghazanfari et al. (2006)
	S. thymbra	Karabay-Yavasoglu et al. (2006)
	Herbicidal activity	S. montana	Espaillat et al. (1993) and Grosso et al. (2010)
S. thymbra		Qasem (2002)
S. thymbrifolia		Qasem (2002)
Effect on muscles		S. obovata subsp. obovata var. valentina	Derojas et al. (1994)
	S. obovata subsp.  obovata var. obovata	Derojas et al. (1994)
	S. viminea	Suarez et al. (2003)
	Butrylcholinesterase inhibitory	S. cuneifolia	Orhan et al. (2008)
S. parvifolia		Cabana et al. (2013)
Aflatoxin B1 production inhibitory activity		S. khuzestanica	Gorran et al. (2013)
	S. macrosiphonia	Gorran et al. (2013)
	Hepatoprotective activity	S. khuzestanica	Ahmadvand et al. (2012), Basiri et al. (2007), Khosravinia et al. (2013), and Saadat et al. (2004)
S. macrostema		Gutierrez (2013) and Gutierrez and Navarro (2010)
Nephroprotective activity		S. khuzestanica	Khosravinia et al. (2013) and Tavafi et al. (2011)
	S. macrostema	Gutierrez (2013)
	Effect of immune system	S. bachtiarica	Pirbalouti et al. (2012)
S. hortensis		Stef et al. (2012)
Effect on productive performance		S. hortensis	Stef et al. (2012)
	S. khuzestanica	Haeri et al. (2006), Rezvanfar et al. (2008a, 2008b), and Safarnavadeh and Rastegarpanah (2011)
	Acaricidal activity	S. hortensis	Aslan et al. (2004)
S. thymbra		Cetin et al. (2010)
Antiproliferative / Antitumor activities		S. boliviana	Miao et al. (1998)
	S. montana	Cetojevic-Simin et al. (2008) and Lampronti et al. (2006)
	Antidiarrheal activity	S. hortensis	Hajhashemi et al. (2000)
S. montana		Skocibusic and Bezic (2003)
Relaxant effect (antispasmoidal activity)		S. hortensis	Boskabady et al. (2007) and Hajhashemi et al. (2000)
	S. parvifolia	Debenedetti et al. (2002) and Hnatyszyn et al. (2003)
	Antigenotoxic activity	S. hortensis	Behravan et al. (2006) and Mosaffa et al. (2006)
Antihepatoma activity	S. hortensis	Lin et al. (2002)
Antilipase activity	S. laevigata	Villa-Ruano et al. (2013)
Neuroprotective activity	S. khuzestanica	Kaeidi et al. (2013)
Nematocidal activity	S. montana	Barbosa et al. (2010, 2012)
Hypolipidemic / Hypoglycemic activities	S. khuzestanica	Ahmadvand et al. (2012), Sarkhail and Rahmani (2011)
Antiplasmoidal activity	S. thymbra	Dell'Agli et al. (2012)
Amoebicidal activity	S. cuneifolia	Malatyali et al. (2012)
Scolicidal activity (Effect on hydatic cysts)	S. khuzestanica	Zibaei et al. (2012)
Contact toxicity and persistance	S. hortensis	Heydarzade and Moravvej (2012)
Photoactivity	S. montana	Matos (2012)
Effect on peripheral blood temperature	S. montana	Masuda et al. (2011)
Effect on body temperature who experiencing cold sensitivity	S. montana	Masuda et al. (2013)
Effect on serum lipid profile	S. khuzestanica	Ahmadvand (2011)
Effect on human ulcerative colitis	S. khuzestanica	Rastegarpanah et al. (2011)
Effect on premature ejaculation	S. montana	Zavatti et al. (2011)
Effect on vitality and healthiness of cereals	S. hortensis	Sas-Piotrowska and Piotrowski (2011)
Effect on blood sugar and lipid levels	S. khuzestanica	Vosough-Ghanbari et al. (2010)
Molluscicidal activity	S. hortensis	Kozlowski and Kozlowska (2009)
Larvicidal activity	S. hortensis	Pavela (2009)
Ovicidal activity	S. thymbra	Ayvaz et al. (2009)
Effect on blood levels of sex hormones	S. khuzestanica	Rezvanfar et al. (2008b)
Effect on DNA integrity and chromatin quality	S. khuzestanica	Rezvanfar et al. (2008b)
Antidiabetic activity	S. thymbra	Loizzo et al. (2008a)
Effect on central nervous system (CNS)	S. mexicana	Estrada-Reyes et al. (2008)
Antihelmintic activity	S. hortensis	Urban et al. (2008)
Inhibition on blood platelet adhesion, aggregation and secretion	S. hortensis	Yazdanparast and Shahriyary (2008)
Effect on jejenum muscular contractility	S. macrostema	Arroyo et al. (2004)
Management of recurrent aphthous stomatitis	S. khuzestanica	Amanlou et al. (2007)
Effect on weakening	S. khuzestanica	Fallahi et al. (2007)
Trypanocidal activity	S. parvifolia	Sulsen et al. (2006)
Antiprotozoal activity	S. parvifolia	van Baren et al. (2006)
Effect on rhinosinusitis	S. hortensis	Uslu et al. (2003)
Effect on motor coordination	S. viminea	Suarez et al. (2003)
Effect on peristaltic activity	S. viminea	Suarez et al. (2003)
Vasodilatory effect	S. obovata	de Rojas et al. (1996a, 1999)
Vasorelaxant effect	S. obovata	de Rojas et al. (1996b)
Diuretic activity	S. montana subsp. montana	Stanic and Samarzija (1993)
Amyloid beta protein aggregation inhibitory	S. hortensis	Ishigaki et al. (2013)
Matrix metalloproteinase inhibitory activity	S. hortensis	Zeidan-Chulia et al. (2013)

According to literature survey, S. hortensis L. and S. montana L. are the most common species evaluated for their antimicrobial activities. In the majority of studies, Gram-negative and Gram-positive bacteria were tested together with fungi. In several studies, only antifungal activities of these species were reported.

As can be seen in the Phytochemistry section of this paper, volatiles of many Satureja species are mainly characterized with their oxygenated monoterpenes (mainly thymol and carvacrol). According to majority of reports, especially presence and amount of carvacrol is the main indicator of antimicrobial activity. As expected, plant species which have high thymol and carvacrol exhibited strong antimicrobial activity.

In most of the prior studies reported before, the year 2000 points that antimicrobial activity was determined by using traditional qualitative techniques such as agar-well and/or agar-disc diffusion. Later 2000, minimum inhibitory concentration (MIC) and minimum bactericidal concentration (MBC) tests were, in general, accompanied to these techniques. In recent years, usages of new techniques such as biodegradable composite biofilms coated with the target material have become widespread. Shojaee-Aliabadi et al. (2013) have characterized biodegradable composite kappa-carrageenan films incorporated with S. hortensis in terms of their physical, optical, mechanical, barrier, and antioxidant properties in addition to the antimicrobial effectiveness of films contrary to five pathogens. Films with the oil effectively inhibited micro-organisms were tested. The authors claimed that S. hortensis oil, as a natural antibacterial agent, can potentially be used in packaging in a wide range of food products, particularly those are highly oxidative and microbial sensitive.

Discussion for the antimicrobial activity potential of Satureja species can be extended. However, due to the page limitations, it should be the subject of another paper.

Antioxidant activity

Reported studies on antioxidant activity potential of Satureja are presented in Table 1. As can be seen from the table, 23 different plant species have been reported for their potential activity in between 1945 and 2014.

In the majority of studies, at least two or more techniques were used together in order to determine the antioxidant activity from different aspects. According to literature data, β-carotene bleaching assay was found as the most commonly used antioxidant test system. Determination of the free radical scavenging potential is another commonly used method. Among the free radicals; DPPH, ABTS, TBARS, superoxide, and nitric oxide are the other most frequently tested parameters.

According to these reports, essential oils of S. hortensis, S. khuzestanica Jamzad, and S. montana, in general, found to have great antioxidant activity due to their oxygenated monoterpenes (especially, carvacrol and thymol contents). However, polar phytochemicals are underlined as biologically active compounds responsible from this activity too. Due to their great antioxidant potential activity, Sature species are regarded as reliable sources for consumption. Their importance, especially, in food industry and ethnopharmacology has strictly been emphasized in these reports.

Of course, there is a lot more to be said about the antioxidant potential activity of this genus. However, it is not possible to discuss the activities of each species here in detail. In fact, antioxidant potential activity of Satureja species should be the subject of a separate article.

Cytotoxic activity

According to the literature survey, cytotoxic activities of 12 Satureja species (S. atropatana Bung., S. hortensis, S. intermedia CA Mey, S. kitaibelii Wierzb. ex Heuff., S. khuzestanica, S. montana, S. punctata subsp. punctate K. Schum., S. sahendica Bornm., S. spicigera (C. Koch) Boiss., and S. thymbra L.) have been evaluated by several researchers (Table 1). By these studies, cytotoxicity of the essential oils and/or extracts of these species was evaluated on several cell lines such as J774 macrophage, 5637, KYSE, Fem-X Human Malignant Melanoma, Vero, SW480, MCF7, JET3, A549, THP-1, and HT29/219.

Cytotoxic activity of S. hortensis on J774 macrophage cell line was tested by Moradi et al. (2008). According to this report, it was toxic at higher concentrations than those needed to inhibit the parasite cell growth (IC₅₀ value = 45.13 μM).

Satureja intermedia is another species in which cytotoxic activity was evaluated by Sadeghi et al. (2013). According to data presented in this report, the IC₅₀ value of the sample against 5637 and KYSE cell lines was found to be 156 μg/mL.

Stanojkovic et al. (2013) evaluated the cytotoxic activity of the methanol extract of S. kitaibelii. The sample exhibited strong activity against Fem-X Human Malignant Melanoma cells (IC₅₀ value = 39.66 μg/mL) and moderate activity against other cancer cell lines (IC₅₀ value from 138.06 μg/mL against estrogen-dependant breast cancer cell line MDA-MB-361–173.15 μg/mL against a human epithelial cervical cancer cells HeLa).

Another plant species investigated fot its cytotoxic activity is S. khuzestanica (Yousefzadi et al., 2014). Essential oil of this plant significantly reduced cell viability of Vero, SW480, MCF7, and JET 3 cells in a dose-dependent manner; IC₅₀ values calculated for each cell types were 31.2, 62.5, 125, and 125 mg/mL.

Toxicity potential of S. punctata subsp. punctata on human monocytic leukemia cells (THP-1) and erythrocytes was evaluated by Tariku et al. (2010). According to this report, the sample indicated high toxicity on THP-1 cells (CC₅₀ value = 0.013–350 nL/mL with selectivity index between 0.001 and 28) and erythrocytes (with LC₅₀ value 0.35–1.52 μg/mL).

Satureja sahendica is another species being investigated for its cytotoxic activity. Essential oil of this species was tested by Yousefzadi et al. (2012). The oil significantly reduced cell viability of MCF7, Vero, SW480, and JET 3 cells in a dose-dependent manner with the IC₅₀ values of 15.6, 125.0, and 250.0 μg/mL, respectively.

Satureja. odora (Gris.) Epl. and S. parvifolia (Phil.) Epl. were also evaluated by Mongelli et al. (1996). According to this report, the toxicity of the dichloromethane extract of these species on brine shrimp was tested and a promising activity was observed (LC₅₀ value <200 μg/mL).

Saab et al. (2012) evaluated the cytotoxicity of S. thymbra by MTT assay in Vero cells. The authors found that S.thymbra exhibited the highest selective index (SI).

As can be seen from the reports presented in this section, most of the studies were carried out with crude extracts and/or essential oils. However, active phytochemicals responsible from cytotoxic activity was only reported by Stanojkovic et al. (2013) and Gohari et al. (2012).

In the first case, rosmarinic acid captured the G2/M phase cell cycle in Fem-X cells, against which both methanol extract and rosmarinic acid possessed the best cytotoxic activity. In the latter case, brine shrimp lethality and four cancerous cell lines HT29/219, Caco (2), NIH-3T3, and T47D were used for cytotoxicity evaluations. 5,7,3′,5′-Tetrahydroxy flavanone was effective against Artemia salina Willd. larva (LC₅₀ value 2.0 μg/mL) and only 5,4′-dihydroxy-3′-methoxyflavanone-7-(6″-O-α-l-rhamnopyranosyl)-β-d-glucopyranoside demonstrated an IC₅₀ value of 98.7 μg/mL on T47D (human, breast, and ductal carcinoma).

Insecticidal activity/insect repellant activity/fumigant toxicity

As far as the literature survey could designate, insecticidal activity of the members of Satureja has been investigated since 1993. Within these reports; S. hortensis, S. thymbra, S. spinosa L., S. parnassica subsp. parnassica Heldr. Sart. ex Boiss., and S. montana have proven to have insecticidal activities. Most of these studies have focused on S. hortensis. It is followed by S. thymbra. In the majority of studies, essential oils were used as target agents. In many reports, essential oils of species are shown to have stronger effect on insects than those of extracts.

Insecticidal activity potential of the members of Satureja is presented in Table 1. As far as the literature survey could designate, plant species have been evaluated against 21 different insect species. Ephestia kuehniella Zeller is the most commonly tested organism (Karaborklu et al., 2011; Maedeh et al., 2011; Sarac & Tunc, 1995a). Ephestia kuehniella is followed by Culex quinquefasciatus (Say) (Pavela, 2008, 2009; Pavela et al., 2008). Another member of Culex (C. pipiens biotype molestus L.) was also evaluated for its sensitivity against the essential oils of three different Satureja species (S. spinosa, S. parnassica subsp. parnassica, S. thymbra, and S. montana) (Michaelakis et al., 2007). Toxicity of the essential oil vapour of S. hortensis was also evaluated against cotton whitefly, Bemisia tabaci (Genn.) (Homoptera: Aleyrodidae) and cowpea seed beetle, Callosobruchus maulatus (F.) (Aslan et al., 2004; Ebadollahi et al., 2012; Heydarzade & Moravvej, 2012; Zandi-Sohani, 2011).

Leptinotarsa decemlineata (Say.), Plodia interpunctella (Hübner.), Acanthoscelides obtectus (Say.), Bruchus dentipes (Baudi), Camptomyia corticalis (H. Loew), Drosophila melanogaster (Meig), Hyalomma marginatum (Koch), Lipaphis pseudobrassicae (Dav.), Musca domestica (L.), Sitophilus granaries (L.), S. oryzae (L.), Spodoptera littoralis (Boisduval), Tetranychus urticae (Koch), Tribolium castaneum (Herbst), and T. confusum (Jacquelin du Val.) are the other organisms of which sensitivities were tested in the presence of Satureja species (Aslan et al., 2004; Cetin et al., 2010; Ebadollahi et al., 2012; Heydarzade & Moravvej, 2012; Karpouhtsis et al., 1998; Kim et al., 2012; Maedeh et al., 2011; Pavela et al., 2008, 2009; ; Sarac & Tunc, 1995a; Tozlu et al., 2011; Yildirim et al., 2011).

In addition to insecticidal activity, insect repellant properties of S. hortensis, S. khuzestanica, S. montana, S. parvifolia, and S. thymbra have also been investigated.

Nottingham and Hardie (1993) and Nottingham et al. (1991) investigated the insect-repellant activity of S. hortensis. In the first report, Brevicoryne brassicae (L.) and Aphis fabae (L.) were subjected to odor of S. hortensis in an olfactometer. Aphis fabae was repelled by plant, while no effect was observed on B. brassicae (Nottingham & Hardie, 1993). According to another study reported by the same research group, volatile of the same plant repelled B. brassicae and A. fabae effectively (Nottingham et al., 1991).

Insect repellant activity of S. khuzestanica was evaluated by Kayedi et al. (2011, 2014). According to the 2014 report, separated solutions with 10%, 20%, and 40% concentrations of essential oil in 99.6% ethanol were prepared. The concentration of repellant was found to be in association with the number of Arthropods bites. The oil exhibited repellency effect even with 10% concentration (Kayedi et al., 2014).

Picard et al. (2012) investigated the repellency effect of S. montana against Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae) adult females. According to this report, the oil at 0.5% concentration showed great repellency effect.

Insect-repellant activity of S. parvifolia was also reported by Lima et al. (2011). According to this study, the oil showed moderate repellent property against Triatoma infestans (Klug) which is the Chagas disease vector.

Finally in this section, repellant activity of S. thymbra will be discussed. According to the literature survey, the activity of this plant species was investigated by Sarac and Tunc (1995a). The oil repelled Sitophilus oryzae adults in food preference tests. In the presence of oil, 5.2–17.4% of the test insects settled in the treated food. According to this report, the oil gave repellency falling into the range of promising repellents, that is, presumably, 40.1–60% repellency as an overall average of an 8 weeks exposure.

Acetylcholinesterase (AChE) and butyrylcholinesterase (BChE) inhibitory activities

As it is well known, AChE and butyrylcholinesterase (BChE) are the chief enzymes of Alzheimer's disease. Inhibitory activities of S. cuneifolia (Ten.), S. khuzestanica, S. montana, S. parvifolia, and S. thymbra were evaluated for their AChE and BChE.

Orhan et al. (2008) tested AChE and BChE inhibitory activities of the essential oil obtained from S. cuneifolia. Essential oil showed a very high inhibitory activity (over 80%) against AChE.

AChE inhibitory activity of S. khuzestanica was also tested by Basiri et al. (2007). In this study, the protective effect of this sample on toxicity of Malathion, a commonly used organophosphorus (OP), was investigated. Malathion inhibited erythrocyte AChE and increased hepatic cells GP and PEPCK activities. On the contrary, co-administration of essential oil resulted in restoration of malathion-induced changes in hepatic cells GP and PEPCK activities and levels of blood AChE and glucose. According to this report, it is concluded that the oil interferes with malathion-induced stimulation of hepatic cells glycogenolysis and gluconeogenesis through its antioxidant potential and increasing AChE activity.

AChE inhibitory activity of the essential oil of S. montana was investigated by Silva et al. (2009). According to this study, inhibitory activity over AChE by S. montana was assessed as a potential indicator for the control of Alzheimer's disease. The supercritical non-volatile fractions, which showed the highest content of (+)-catechin, chlorogenic, vanillic, and protocatechuic acids, also inhibited selectively and significantly BChE.

Another plant species evaluated for its AChE and BChE inhibitory activities is S. parvifolia. According to a study reported by Cabana et al. (2013), three different polar extracts (decoction, ethanol, and hydrolate) of this species were studied. All extracts showed excellent AChE and BChE inhibitory capacities.

Satureja thymbra was also evaluated for its cholinesterase inhibitory effects against AChE by Ozturk (2012). According to this study, a promising enzyme inhibition activity was obtained in the presence of essential oil.

Antiviral activity

According to the literature survey, antiviral activities of S. boliviana Briq., S. cuneifolia, S. hortensis, S. montana, and S. thymbra have been reported against several types of viruses including Herpes simplex type-1 (HSV-1), Parainfluenza type-3 (PI-3), Poliovirus type 1, Human immunodeficiency virus type-1 (HIV-1), Vesicular stomatitis virus (VSV), Cucumber mosaic virus (CMV), and SARS-CoV (Table 1).

Antiviral activity potential of S. thymbra was investigated by two different research groups (Loizzo et al., 2008b; Saab et al., 2012). On one hand, Saab et al. (2012) examined the in vitro antiviral activity of S. thymbra against HSV-1 infection on monkey kidney cells. According to this report, the sample showed a high selective index (SI) and is, therefore, potentially be used for treatment of HSV-1 disease. On the other hand, according to another study reported by Loizzo et al. (2008b), S. thymbra essential oil was evaluated for its inhibitory activity against SARS-CoV and HSV-1 replication in vitro by visually scoring of the virus-induced cytopathogenic effect post-infection. The oil exerted moderate activity against the both viruses.

Ethanolic and aqueous extracts of S. boliviana were tested for their antiviral activities against HSV-1, VSV, and poliovirus type 1 by Abad et al. (1999). The aqueous extract showed remarkable antiviral activity against HSV-1 and VSV.

The essential oil of S. cuneifolia, as well as the widely encountered components in essential oil was screened for their antiviral activities by Orhan et al. (2012) against HSV-1 and PI-3. The oil and compounds exhibited strong antiviral effects against HSV-1, ranging between 0.8 and 0.025 μg/mL. The samples tested were less effective against PI-3, with results ranging between 1.6 and 0.2 μg/mL.

The anti-HIV-1 activity of S. montana was also evaluated by Yamasaki et al. (1998). Especially, aqueous extract showed a potent anti-HIV-1 activity (with an ED value of 16 μg/mL). Active components in the extract were found to be water-soluble polar substances. Additionally, the aqueous extract inhibited giant cell formation in co-culture of Molt-4 cells with and without HIV-1 infection and showed inhibitory activity against HIV-1 reverse transcriptase.

The essential oil of S. montana (L.) ssp. variegata [(Host) P. W. Ball] was applied on local hosts Chenopodium amaranticolor (Coste & Reyn.) and C. quinoa (Willd.) simultaneously with the infecting viruses by Dunkic et al. (2010). According to this report, the number of local lesions on both TMV and CMV infected plants was reduced for 29.2% and 24.1%, respectively. When applied individually for each virus, thymol was more effective in reducing CMV infection (33.2%), while carvacrol was more effective in reducing the TMV infection (34.3%).

Antinociceptive/analgesic activity

Satureja cuneifolia, S. hortensis, S. khuzestanica, S. thymbra, and S. viminea (Burm.) are the plant species of which antinociceptive/analgesic activities were reported elsewhere (Table 1).

Among these species, the activity of S. hortensis was studied by Hajhashemi et al. (2002, 2012). In the first case, hydroalcoholic and polyphenolic extracts and essential oil of S. hortensis were evaluated. According to this report, hydroalcoholic extract significantly reduced pain responses in early and late phases of the formalin test whereas the polyphenolic extract and essential oil were only effective in the late phase of the formalin test (Hajhashemi et al., 2002). In the second case, the same extracts were evaluated for their analgesic activities using light tail flick, formalin, and acetic acid-induced writhing tests in mice. According to the results, in the light tail flick test, neither the essential oil nor the extracts exerted any significant effect. In the formalin test, hydroalcoholic extract (500–2000 mg/kg, p.o.), polyphenolic fraction (250–1000 mg/kg, p.o.), and the essential oil (50–200 mg/kg, p.o.) showed analgesic activity. According to the authors, involvement of opioid and adenosine receptors mediates the antinociception (Hajhashemi et al., 2012).

In a study reported by Aydin et al. (1996), essential oil of S. cuneifolia was investigated for its analgesic activity by the using tail-flick method in mice. According to the findings obtained from this study, analgesic activity is strictly related to the carvacrol content of essential oil.

Satureja khuzestanica has been used as analgesic and antiseptic agents among the inhabitants of southern parts of Iran. Amanlou et al. (2005) investigated antinociceptive effect of the hydroalcoholic extract of this plant using the carrageenan-induced rat paw edema and formalin tests. The extract showed antinociceptive activity in a dose-dependent manner (10–150 mg/kg; i.p.) at the second phase of formalin test which was comparable with morphine (3 mg/kg; i.p.). According to authors, the presence of flavonoids, steroids, essential oil, mainly carvacrol, and tannin might be responsible for antinociceptive activity of this plant.

Antinociceptive activity of the essential oil of S. thymbra was also measured by the formalin test in mice and by the light tail-flick and hot-plate methods in rats by Karabay-Yavasoglu et al. (2006). According to the authors, although the essential oil produced an antinociceptive effect during both the early (50 and 100 mg/kg) and late phases (25, 50, and 100 mg/kg) of the formalin test, it did not exert any significant antinociceptive effect in tail-flick and paw-edema tests, respectively. Therefore, it may have central analgesic activity in mice and rats.

Finally, in this section, analgesic activity of the essential oil of S. viminea will be discussed. According to a study reported by Suarez et al. (2003), a very clear and significant analgesic effect was observed with the oral administration of the oil (1000 mg/kg). The LD₅₀ value of the oil was determined as 556.8 mg/kg.

Antileishmanial activity

According to the literature survey, antileishmanial activities of S. bachtiarica Bunge, S. hortensis, S. khuzestanica, and S. punctata subsp. punctata have previously been reported several times (Table 1).

Antileishmanial activity of S. khuzestanica was studied by Sadeghi-Nejad et al. (2011) and Kheirandish et al. (2011), independently. Sadeghi-Nejad et al. (2011) assessed antiprotozoal activity of ethanolic and methanolic extracts of the leaves of S. khuzestanica against Leishmania major promastigotes. The extracts inhibited parasite after 24 h incubation against L. major promastigotes, which gave the IC₁₀₀ values of 2.4 and 4.8 mg/mL and IC₅₀ values of 0.3 and 0.6 mg/mL, respectively. According to the authors, leaf extracts contain active compounds, which could serve as an alternative agent in the control of cutaneous leishmaniasis. However, further studies would, therefore, be needed to see its in vivo clinical response and associated toxicities.

Antileishmanial activity of the essential oil of S. khuzestanica was also reported in animal model of leishmaniasis by Kheirandish et al. (2011). According to this report, the results indicated that the oil has an effect on preventing death in infected mice. According to the results obtained by the authors, it seems that the oil is not only antifungal, antiviral, and antibacterial but also an antiparasitic agent.

Antileishmanial activity of the essential oil of S. bachtiarica was evaluated by Mohammadpour et al. (2012). According to this report, the oil showed higher activity against L. major than the standard antileishmanial drug, glucantime, and all the parasites were killed after 24 h.

Antileishmanial activity of the ethanolic extract of S. hortensis was assessed against L. major promastigotes by Manjili et al. (2012). Satureja hortensis exhibited high antileishmanial activity (IC₅₀ value 15.625 μM). According to the authors, S. hortensis contains active compounds, which could serve as alternative agents in the control of cutaneous leishmaniasis.

Essential oil of S. punctata ssp. punctata from Ethiopia was also screened for its leishmanicidal activity against promastigote and axenic amastigotes of L. donovani and L. aethiopica by Tariku et al. (2010). The oil showed effect on promastigotes (MIC value 76.5–312.5 nL/mL) and amastigotes (EC₅₀ value 4.06–131.00 nL/mL) of L. donovani and L. aethiopica.

Genotoxic activity

Studies carried out on genotoxic activity potentials of the members of Satureja are presented in Table 1. As can be seen from the table, there are limited number of studies under this topic. Plant species evaluated for their genotoxic effect are S. hortensis, S. montana, and S. thymbra.

Genotoxic properties of the essential oils of S. hortensis and S. montana were studied with Bacillus subtilis rec-assay and Salmonella/microsome reversion assay by Zani et al. (1991). Both oils exhibited moderate activity in rec-assay and Salmonella tests. In this report, advantages of the combined use of these two short-term microbial assays in genotoxic studies were also discussed.

The essential oil and its main constituents, carvacrol and thymol, of S. thymbra were also tested for their genotoxic activities on Drosophila by Karpouhtsis et al. (1998). According to this report, the somatic mutation and recombination test on Drosophila revealed that, among the five compounds studied, only thymol exhibited genotoxic activity.

Anti-inflammatory activity

Satureja hortensis, S. khuzestanica, and S. thymbra are the plant species of which anti-inflammatory activities were discussed by Hajhashemi et al. (2002, 2012), Ghazanfari et al. (2006), Amanlou et al. (2005), and Karabay-Yavasoglu et al. (2006) (Table 1).

Anti-inflammatory activity of S. hortensis was reported by the same research group at different times (Hajhashemi et al., 2002, 2012). According to the first report, hydroalcoholic and polyphenolic extracts and essential oil of S. hortensis seeds were tested for their activity potentials. For the evaluation of anti-inflammatory activity, the carrageenan-induced rat paw edema test was used. All three fractions were found to reduce paw edema in the carrageenan test. According to the authors, the results of this study substantiated the traditional use of S. hortensis plant seeds in inflammatory ailments (Hajhashemi et al., 2012). In the second case, anti-inflammatory effects of the same fractions of S. hortensis were assessed using carrageenan-induced paw edema test in rats. According to this report, polyphenolic fraction (1000 mg/kg, p.o.) and the essential oil (200 mg/kg) reduced edema caused by carrageenan (Hajhashemi et al., 2002).

The essential oil from S. khuzestanica was evaluated for its activity against inflammatory bowel disease (IBD) by Ghazanfari et al. (2006). According to this report, the beneficial effect of essential oil (1500 ppm) was found comparable with that of prednisolone. The authors claimed that anti-inflammatory potential of this species may be the mechanisms by which this plant protects animals against experimentally induced IBD.

Anti-inflammatory activity of the same plant species was also investigated by Amanlou et al. (2005). According to this report, a similar anti-inflammatory activity was observed between S. khuzestanica hydroalcoholic extract (150 mg/kg; i.p.) and indomethacin (4 mg/kg; i.p.) in the carrageenan test. The authors claimed that the presence of flavonoids, steroids, essential oil, mainly carvacrol and tannin, might be responsible for anti-inflammatory activity of this plant.

The last plant species of which anti-inflammatory activity reported is S. thymbra. Karabay-Yavasoglu et al. (2006) reported the activity of the essential oil of this plant. According to the authors, the oil did not exert any anti-inflammatory effect in mice and rats.

Herbicidal activity

As far as the literature survey could designate, herbicidal activities of S. montana, S. thymbra, and S. thymbrifolia Hedge & Feinbrun. have previously been reported (Table 1).

Qasem (2002) has studied the possible role of allelopathy in controlling branched broomrape (Orobanche ramose L.), through screening different plant species. According to this report, S. thymbra and S. thymbrifolia encouraged and increased parasite infestation. The authors underlined the importance of certain plants as trapping or catching species for the parasite and the potential effects of their root exudates and/or residues on O. ramosa.

The effect of the essential oil of S. montana on seed germination, root and shoot growth of the surviving seedlings of four crops [Zea mays (L.), Triticum durum (Dest.), Pisum sativum (L.), and Lactuca sativa (L.)] and two weeds [Portulaca oleracea (L.) and Vicia sativa (L.)], was investigated by Grosso et al. (2010). The oil affected both by crop and weeds and can be appropriate for uncultivated fields.

The activity of S. montana was also reported by Espaillat et al. (1993). In this study, norflurazon proved to be the least phytotoxic herbicide. Crop vigor of S. montana was not affected by norflurazon applied at 0.56 kg a.i. ha⁻¹ (active ingredient per hectar) (0.5×). Bentazon at 0.84 kg a.i. ha⁻¹ (0.5×) and dicamba at 0.02 kg a.i. ha⁻¹ (0.5×) showed potential for labeling in the presence of plant sample. According to this report, simazine killed exposed plants at all rates.

Effect on muscles

According to the literature survey, two different varieties of S. obovata Lag. (S. obovata subsp. obovata var. valentina and S. obovata subsp. obovata var. obovata) and S. viminea were evaluated for their effects on muscles (Derojas et al., 1994; Suarez et al., 2003).

Suarez et al. (2003) have tested the aqueous extract and the essential oil of S. viminea. Muscle strength was assessed through a grasping test where mice were hung by their fore-limbs 40 cut above the base on a horizontal metal stainless bar. According to the results, a dose-related spontaneous motor activity reduction was observed. Exploratory behavior and curiosity were also diminished.

According to a study, aqueous extracts of S. obovata subsp. obovata var. valentina and S. obovata subsp. obovata var. obovata exhibited a dose-dependent inhibitory effect on the contractions induced by acetylcholine in rat duodenum and by noradrenaline in rat aorta. The extracts also produced relaxant effects in both tissue preparations precontracted with K⁺ (75 mM). Vasodilatory effect of var. valentina was stronger than the other. The authors underlined the smooth muscle relaxant effects of these extracts.

Other activities

Apart from the activities discussed above, as can be seen in Table 1, approximately 50 different activities of the members of Satureja genus have also been reported at somewhere else in between 1945 and 2013. Among these, the most commonly studied species are S. hortensis and S. khuzestanica.

It is not possible to discuss full biological activity profile of Satureja species here due to the high number of reports under this topic. According to the literature survey, aflatoxin B1 production inhibitory, hepatoprotective, nephroprotective, acaricidal, antiproliferative/antitumor, antidiarrheal, relaxant (antispasmoidal), antigenotoxic, antihepatoma, antilipase, neuroprotective, nematocidal, hypolipidemic/hypoglycemic, antiplasmoidal, scolicidal (effect on hydatic cysts), molluscicidal, larvicidal, ovicidal, antidiabetic, antihelmintic, trypanocidal, antiprotozoal, amyloid beta protein aggregation inhibitory, and matrix metalloproteinase inhibitory activities of Satureja species have been reported so far. In addition to these activities, the effect of Satureja species on immune system, productive performance, contact toxicity and persistance, peripheral blood temperature, body temperature who experiences cold sensitivity, serum lipid profile, human ulcerative colitis, premature ejaculation, vitality and healthiness of cereals, blood sugar and lipid levels, blood levels of sex hormones, DNA integrity and chromatin quality, central nervous system (CNS), blood platelet adhesion, aggregation and secretion, jejenum muscular contractility, weakening, rhinosinusitis, motor coordination, and peristaltic activity have also been identified by many reports. According to the literature survey, antimicrobial, antioxidant, antidiabetes, antihyperlipidemic, and reproductive stimulation performance of Satureja species have also been discussed in a review by Momtaz and Abdollahi (2010).

Phytochemistry

Volatiles (terpenes)

Major volatiles of the members of Satureja are presented in detail in Table 2. As it is well known, Satureja species are especially rich in oxygenated monoterpenes. Among these, the best known are thymol and carvacrol. Characteristic terpenoids of Satureja species are also presented as Figure 1.

Figure 1. Characteristic terpenoids of Satureja species.

Table 2. Major terpenoids of Satureja species.

Major Terpenoids	Molecular formula	Average mass (Da)	Plant Species	Reference
(-)-Cadin-4-en-1-ol	C15H26O	222.366302	S. gilliesii	Labbe et al. (1993)
(+)-Spathulenol	C15H24O	220.350494	S. sahendica	Taherpour et al. (2008)
(+)-T-cadinol	C15H26O	222.366302	S. gilliesii	Labbe et al. (1993)
(E)-Nerolidol	C15H26O	222.366302	S. punctata subsp. punctata	Tariku et al. (2010)
(Z)-β-Ocimene	C10H16	136.233994	S. montana subsp. montana	Slavkovska et al. (2001)
			S. montana subsp. pisidica	Slavkovska et al. (2001)
			S. subspicata	Bezic et al. (2009)
			1,8-Cineole  (syn. eucalyptol)	C10H18O	154.249298	S. atropatana	Rustaiyan et al. (2004)
S. boliviana	Hatami et al. (2011) and Viturro et al. (2000)
S. mutica	Rustaiyan et al. (2004)
S. wiedemanniana	Altundag and Aslim (2011)
Bicyclogermacrene	C15H24	204.351105				S. boliviana	Viturro et al. (2000)
Borneol	C10H18O	154.249298	S. bachtiarica	Hadian et al. (2012), Sefidkon and Jamzad (2000)
			S. cuneifolia	Kan et al. (2006), Mirjana et al. (2004), and Tommasi et al. (2008, 2009)
			S. hortensis	Kubatova et al. (2001)
			S. horvatii	Lakusic et al. (2008)
			S. horvatii subsp. macrophylla	Dardioti et al. (2010)
			S. kitaibelii	Mihajilov-Krstev et al. (2011)
			S. macrosiphonia	Amiri (2011)
			S. montana	Barbosa et al. (2012), Mirjana and Nada (2004), Radonic and Milos (2003b), and Skocibusic and Bezic (2004)
			S. montana subsp. montana	Slavkovska et al. (2001)
			S. montana subsp. pisidica	Slavkovska et al. (2001)
			S. sahendica	Taherpour et al. (2008)
			S. subspicata	Skocibusic et al. (2006)
			S. thymbra	Piras et al. (2011)
			S. visianii	Vidic et al. (2009)
			Bornyl acetate	C12H20O2	196.285995	S. biflora	Vagionas et al. (2007)
Camphene	C10H16	136.233994	S. hortensis	Kizil (2009)
			S. isophylla	Habibi et al. (2007)
			S. kitaibelii	Slavkovska et al. (2001)
			S. montana subsp. montana	Slavkovska et al. (2001)
			S. montana subsp. pisidica	Slavkovska et al. (2001)
			Camphor	C10H16O	152.233398	S. douglasii	Peer and Langenheim (1998)
S. isophylla	Habibi et al. (2007) and Sefidkon and Jamzad (2006)
S. x delpozoi hybrid (S. intricata x S. obovata)	Jordan et al. (2010)
S. visianii	Bezic et al. (2009), Dunkic and Bezic (2008), and Vidic et al. (2009)
Carvacrol	C10H14O	150.21761				S. atropatana	Moradi et al. (2008)
			S. bachtiarica	Ahanjan et al. (2011b), Hadian et al. (2012), Hashemi et al. (2011), Meshkatalsadat and Shabaninejad (2011), Pirbalouti et al. (2013)
			S. boissieri	Kurkcuoglu et al. (2001) and Sajjadi and Baluchi (2002)
			S. capitata	Napoli et al. (2010)
			S. cilicica	Cetojevic-Simin et al. (2012) and Schulz et al. (2005)
			S. coerulea	Cetojevic-Simin et al. (2012)
			S. cuneifolia	Altun and Goren (2007), Azaz et al. (2005), Baydar et al. (2004), Bezic et al. (2001, 2009, Biavati et al. (2004), Habibi et al. (2007), Kosar et al. (2008), Kulisic et al. (2005), Mirjana et al. (2004), Oke et al. (2009a, 2009b), Orhan et al. (2011), Schulz et al. (2005), Tumen et al. (1998b)
			S. hortensis	Abdollahi et al. (2011) 2012, Adiguzel et al. (2007), Alizadeh et al. (2010a), Alizadeh-Salteh et al. (2010b), Annan et al. (2013), Baher et al. (2002), Baser et al. (2004), Dawidowicz and Rado (2010), Deighton et al. (1994), Dikbas et al. (et al. (2010), Dikbas et al. (2011), Djenane et al. (2011a), Ebadi et al. (2011), Esquivel et al. (1999), Figueredo et al. (2012), Ghannadi (2002), Gulluce et al. (2003), Gursoy et al. (2009), Hadian et al. (2010a), Heydarzade and Moravvej (2012), Hoferl et al. (2009), Karami-Osboo et al. (2010), Khajeh (2011), Kizil (2009), Kizil and Uyar (2006), Kizil et al. (2009), Kubatova et al. (2001), Lampronti et al. (2006), Mihajilov-Krstev et al. (2009b), Mumivand et al. (2011), Nedorostova et al. (2011), Novak et al. (2006), Omidbaigi et al. (2007), Pank (2009), Pavela (2009), Pfefferkorn et al. (2008), Pistelli et al. (2013), Razzaghi-Abyaneh et al. (2008), Rezvanpanah et al. (2011), Saharkhiz et al. (2011), Sajfrtova et al. (2013), Sampson et al. (2005), Sefidkon et al. (2006), Shatar and Altantsetseg (2000), Shirzad et al. (2011), Tozlu et al. (2011), Trojakova et al. (2001), and Viturro et al. (1999), Yazici et al. (2011)
			S. horvatii	Lakusic et al. (2008, 2011)
			S. horvatii subsp. horvatii	Dardioti et al. (2010)
			S. icarica	Azaz et al. (2002) and Cetojevic-Simin et al. (2012)
			S. khuzestanica	Farsam et al. (2004), Hadian et al. (2010b, 2011, 2012), Hashemi et al. (2012), Kayedi et al. (2011), Kheirandish et al. (2011), Pirbalouti and Moalem (2013), Pirbalouti et al. (2011c), Saei-Dehkordi et al. (2012), Sefidkon and Ahmadi (2000), and Taban et al. (2013)
			S. laxiflora	Sonboli et al. (2004)
			S. macrantha	Azaz et al. (2005)
			S. mittica	Gohari et al. (2005)
			S. montana	Angelini et al. (2003), Barbosa et al. (2010, 2012, Bezic et al. (2009), Chizzola (2003), Coelho et al. (2012), Damjanovic-Vratnica et al. (2011), de Oliveira et al. (2011, 2012, Djenane et al. (2011b), 2012, Dunkic et al. (2012), Fraternale et al. (2007), Grosso et al. (2009a, 2009b), Haloci et al. (2011), Ibraliu et al. (2011), Kulisic et al. (2005), Marin et al. (2012), Mastelic and Jerkovic (2003), Michaelakis et al. (2007), Miladi et al. (2013), Mirjana and Nada (2004), Nedorostova et al. (2011), Omidbaigi et al. (2007), Palavra et al. (2011), Penalver et al. (2005), Prieto et al. (2007), Radonic and Milos (2003b), Ratajac et al. (2011), Serrano et al. (2011), Sfeir et al. (2013), Skocibusic and Bezic (2003, 2004, Stoilova et al. (2008), and Vidic et al. (2009)
			S. montana subsp. montana	Nhu-Trang et al. (2006)
			S. montana subsp. pisidica	Slavkovska et al. (2001)
			S. montana subsp. variegata	Dunkic et al. (2010)
			S. mutica	Hadian et al. (2012), Sefidkon and Jamzad (2005)
			S. pilosa subsp. origanita	Dardioti et al. (2012)
			S. rechingeri	Hadian et al. (2010b, 2012), Safari et al. (2011), and Sefidkon et al. (2007)
			S. sahendica	Hassanpouraghdam et al. (2009)
			S. spicigera	Eftekhar et al. (2009), Eminagaoglu et al. (2007), Kotan et al. (2010), and Sefidkon and Jamzad (2004)
			S. spinosa	Michaelakis et al. (2007) and Skoula and Grayer (2005)
			S. subspicata	Skocibusic et al. (2006)
			S. subspicata subsp. liburnica	Dunkic et al. (2007)
			S. subspicata subsp. subspicata	Dunkic et al. (2007)
			S. thymbra	Ayvaz et al. (2009), Azaz et al. (2005), Cetin et al. (2010), Dell'Agli et al. (2012), Economou et al. (2011), Figueredo et al. (2012), Fleisher and Fleisher (2005), Giweli et al. (2012), Glamoclija et al. (2006), Goren et al. (2004), Karousou et al. (2005), Karpouhtsis et al. (1998), Lagouri et al. (1993), Loizzo et al. (2008b), Michaelakis et al. (2007), Muller-Riebau et al. (1997), Mullerriebau et al. (1995), Ozturk (2012), Piras et al. (2011), Sampson et al. (2005), Schulz et al. (2005), Skoula and Grayer (2005), Sokovic et al. (2002), and Ugur et al. (2009)
			S. wiedemanniana	Altundag and Aslim (2011), Baser et al. (2001), Sampson et al. (2005)
			Carvacrol methyl ether	C12H20O2	196.285995	S. horvatii	Lakusic et al. (2008)
S. montana	Radonic and Milos (2003b), Serrano et al. (2011), and Skocibusic and Bezic (2004),
Carvone	C10H14O	150.217606				S. atropatana	Rustaiyan et al. (2004)
			S. douglasii	Peer and Langenheim (1998)
			Caryophyllene	C15H24	204.351105	S. boliviana	Hatami et al. (2011)
S. hortensis	Pistelli et al. (2013)
S. thymbra	Economou et al. (2011)
Caryophyllene oxide	C15H24O	220.350494				S. bachtiarica	Hadian et al. (2012) and Hashemi et al. (2011)
			S. icarica	Azaz et al. (2002)
			S. sahendica	Taherpour et al. (2008)
			S. thymbra	Fleisher and Fleisher (2005)
			cis-3-Hexane-1-ol	C6H14O	102.174797	S. montana	Mastelic and Jerkovic (2003)
cis-Jasmone	C11H16O	164.244095	S. bachtiarica	Ahanjan et al. (2011b) and Meshkatalsadat and Shabaninejad (2011)
cis-Linalool oxide	C10H18O2	170.248703	S. biflora	Matasyoh et al. (2007)
cis-Ocimene	C10H16	136.233994	S. cuneifolia	Biavati et al. (2004)
cis-Piperitenone epoxide	-	-	S. parvifolia	Lima et al. (2011) and Luna et al. (2008)
cis-Piperitone oxide	C10H16O2	168.232803	S. masukensis	Vagionas et al. (2007)
cis-Terpinene	C10H16	136.233994	S. hortensis	Mumivand et al. (2011)
Citronellol	C10H20O	156.265198	S. glabrata	Malagon et al. (2003)
			S. hortensis	Soran et al. (2011)
			Citronellyl acetate	C12H22O2	198.301895	S. glabrata	Malagon et al. (2003)
Cymol	C10H14	134.218201	S. hortensis	Kizil et al. (2009)
Dihydrothymoquinone	C10H12O2	164.201096	S. hortensis	Taborsky et al. (2012)
			S. montana	Taborsky et al. (2012)
			Elemol	C15H26O	222.366302	S. isophylla	Habibi et al. (2007)
endo-1-Bourbonanol	C15H24O	220.350494	S. biflora	Matasyoh et al. (2007)
Eugenol	C10H12O2	164.201096	S. khuzestanica	Farsam et al. (2004)
			S. montana	Mastelic and Jerkovic (2003)
			Farnesol	C15H26O	222.366302	S. montana	Djenane et al. (2012)
Geranial	C10H16O	152.233398	S. forbesii	Ortet et al. (2009)
			S. isophylla	Sefidkon and Jamzad (2006)
			S. punctata	Chagonda and Chalchat (2005)
			S. punctata subsp. punctata	Tariku et al. (2010)
			Geraniol	C10H18O	154.249298	S. hortensis	Soran et al. (2011)
S. laxiflora	Sonboli et al. (2004)
S. montana	Cavar et al. (2008), Mastelic and Jerkovic (2003), Mirjana and Nada (2004), and Radonic and Milos (2003b)
S. robusta	Tchoumbougnang et al. (2009)
Geranyl acetate	C12H20O2	196.285995				S. bachtiarica	Ahanjan et al. (2011b)
			S. laxiflora	Sonboli et al. (2004)
			S. montana	Mirjana and Nada (2004)
			S. subspicata	Skocibusic et al. (2006)
			S. visianii	Bezic et al. (2009)
			Geranyl acetone	C14H24O	208.339798	S. bachtiarica	Meshkatalsadat and Shabaninejad (2011)
Germacrene D	C15H24	–	S. biflora	Matasyoh et al. (2007)
			S. boliviana	Viturro et al. (2000)
			S. corulea	Tumen et al. (1998a)
			S. robusta	Tchoumbougnang et al. (2009)
			Isocaryophyllene	C15H24	204.351105	S. glabrata	Malagon et al. (2003)
Isomenthone	C10H18O	154.249298	S. abyssinica subsp. abyssinica	Tolossa et al. (2007)
			S. boliviana	Dambolena et al. (2009), Hatami et al. (2011), and Senatore et al. (1998)
			S. brevicalix	Senatore et al. (1998)
			S. brownei	Rojas and Usubillaga (2000)
			S. douglasii	Peer and Langenheim (1998)
			S. fruticosa	Coelho et al. (2007, 2012)
			S. multiflora	Niemeyer (2010)
			S. pseudosimensis	Vagionas et al. (2007)
			Isopulegone	C10H16O	152.233398	S. brownei	Rojas and Usubillaga (2000)
Limonene	C10H16	136.233994	S. cuneifolia	Mirjana et al. (2004) and Skocibusic and Bezic (2004)
			S. hortensis	Soran et al. (2011)
			S. kitaibelii	Mihajilov-Krstev et al. (2011) and Slavkovska et al. (2001)
			S. macrantha	Sefidkon and Jamzad (2005)
			S. montana	Bezic et al. (2009)
			S. montana subsp. montana	Slavkovska et al. (2001)
			S. montana subsp. kitaibelii	Konakchiev and Tsankova (2002)
			S. odora	Fuselli et al. (2008)
			S. sahendica	Hassanpouraghdam et al. (2009)
			S. subspicata	Bezic et al. (2009) and Skocibusic et al. (2006)
			S. visianii	Bezic et al. (2009) and Dunkic and Bezic (2008)
			Linalool	C10H18O	154.249298	S. bachtiarica	Hadian et al. (2012)
S. biflora	Matasyoh et al. (2007) and Vagionas et al. (2007)
S. boliviana	Viturro et al. (2000)
S. cuneifolia	Bezic et al. (2001, 2009, Habibi et al. (2007), Kan et al. (2006), Mirjana et al. (2004), Orhan et al. (2011), Savikin et al. (2010), and Tommasi et al. (2008, 2009)
S. hortensis	Kubatova et al. (2001) and Soran et al. (2011)
S. horvatii	Lakusic et al. (2011)
S. horvatii subsp. macrophylla	Dardioti et al. (2010)
S. kitaibelii	Slavkovska et al. (2001)
S. masukensis	Vagionas et al. (2007)
S. montana	Chizzola (2003), de Oliveira et al. (2011), Fraternale et al. (2007), and Ratajac et al. (2011)
S. montana subsp. montana	Slavkovska et al. (2001)
S. montana subsp. pisidica	Slavkovska et al. (2001)
S. parnassica subsp. parnassica	Tzakou and Skaltsa (2003)
S. pilosa	Ozek et al. (2010)
S. pilosa subsp. origanita	Dardioti et al. (2012)
S. punctata	Tchoumbougnang et al. (2009)
S. sahendica	Taherpour et al. (2008)
S. spinosa	Aslan et al. (2004) and Ozek et al. (2010)
S. subspicata	Skocibusic et al. (2006)
Menthol	C10H20O	156.265198				S. atropatana	Rustaiyan et al. (2004)
			S. mutica	Rustaiyan et al. (2004)
			Menthone	C10H18O	154.249298	S. atropatana	Rustaiyan et al. (2004)
S. boliviana	Senatore et al. (1998)
S. brevicalix	Senatore et al. (1998)
S. brownei	Rojas and Usubillaga (2000)
S. odora	Fuselli et al. (2008)
Methyl carvacrol	C11H16O	164.244095				S. montana	Mirjana and Nada (2004)
Methyl chavicol	C14H22O	206.323898	S. atropatana	Rustaiyan et al. (2004)
Methyl thymol	C11H16O	164.244095	S. mittica	Gohari et al. (2005)
			S. montana	Mirjana and Nada (2004)
			mid-Spathulenol	C15H24O	220.350494	S. atropatana	Gohari et al. (2005)
Myrcene (syn. β-Myrcene)	C10H16	136.233994	S. cuneifolia	Altun and Goren (2007)
			S. hortensis	Kizil (2009)
			S. horvatii	Lakusic et al. (2008)
			S. khuzestanica	Farsam et al. (2004) and Taban et al. (2013)
			S. kitaibelii	Slavkovska et al. (2001)
			S. montana	Sfeir et al. (2013)
			S. montana subsp. montana	Slavkovska et al. (2001)
			S. montana subsp. montana	Slavkovska et al. (2001)
			S. sahendica	Hassanpouraghdam et al. (2009)
			S. subspicata	Skocibusic et al. (2006)
			Neral	C10H16O	152.233398	S. forbesii	Ortet et al. (2009)
S. punctata	Chagonda and Chalchat (2005)
S. punctata subsp. punctata	Tariku et al. (2010)
o-Ocimene	C10H16	136.233994				S. cuneifolia	Orhan et al. (2011)
			S. hortensis	Mumivand et al. (2011) and Pistelli et al. (2013)
			S. sahendica	Hassanpouraghdam et al. (2009)
			p-Bisabolene	C15H24	204.351105	S. montana	Grosso et al. (2009a)
p-Cymen-8-ol	C10H14O	150.217606	S. montana subsp. montana	Slavkovska et al. (2001)
p-Cymene	C10H14	134.218201	S. aintabensis	Azaz et al. (2005) and Sampson et al. (2005)
			S. atropatana	Gohari et al. (2005)
			S. bachtiarica	Ahanjan et al. (2011a), Meshkatalsadat and Shabaninejad (2011), Mohammadpour et al. (2012), Pirbalouti et al. (2013), and Sefidkon and Jamzad (2000)
			S. boissieri	Kurkcuoglu et al. (2001) and Sajjadi and Baluchi (2002)
			S. boliviana	Hatami et al. (2011)
			S. cilicica	Schulz et al. (2005)
			S. cuneifolia	Altun and Goren (2007), Bezic et al. (2001), Bezic et al. (2009), Biavati et al. (2004), Eminagaoglu et al. (2007), Kan et al. (2006), Kosar et al. (2008), Mirjana et al. (2004), Oke et al. (2009a,2009b), and Schulz et al. (2005)
			S. edmondi	Masumeh (2006), Sefidkon and Jamzad (2006)
			S. hortensis	Adiguzel et al. (2007), Alizadeh et al. (2010a), Alizadeh-Salteh et al. (2010b), Dawidowicz and Rado (2010), Dikbas et al. (2011), Figueredo et al. (2012), Ghannadi (2002), Gulluce et al. (2003), Gursoy et al. (2009), Hadian et al. (2010a), Heydarzade and Moravvej (2012), Hoferl et al. (2009), Mihajilov-Krstev et al. (2009b), Nedorostova et al. (2011), Pfefferkorn et al. (2008), Pistelli et al. (2013), Sajfrtova et al. (2013), Tozlu et al. (2011), and Viturro et al. (1999)
			S. horvatii	Lakusic et al. (2008)
			S. intermedia	Sefidkon and Jamzad (2005)
			S. khuzestanica	Farsam et al. (2004), Hadian et al. (2012), Sefidkon and Ahmadi (2000)
			S. kitaibelii	Kundakovic et al. (2011), Slavkovska et al. (2001)
			S. laxiflora	Sonboli et al. (2004)
			S. macrantha	Sefidkon and Jamzad (2005)
			S. mittica	Gohari et al. (2005)
			S. montana	Barbosa et al. (2010, 2012), Bezbradica et al. (2005), Bezic et al. (2009), Chizzola (2003), Coelho et al. (2012), Damjanovic-Vratnica et al. (2011), de Oliveira et al. (2011, 2012, Djenane et al. (2012), Fraternale et al. (2007), Grosso et al. (2009a), Ibraliu et al. (2011), Marin et al. (2012), Mastelic and Jerkovic (2003), Michaelakis et al. (2007), Mihajilov-Krstev et al. (2011), Miladi et al. (2013), Mirjana and Nada (2004), Nedorostova et al. (2011), Prieto et al. (2007), Radonic and Milos (2003b_), Ratajac et al. (2011), Sfeir et al. (2013), Skocibusic and Bezic (2004), and Stoilova et al. (2008)
			S. montana subsp. montana	Nhu-Trang et al. (2006), Slavkovska et al. (2001)
			S. montana subsp. pisidica	Slavkovska et al. (2001)
			S. mutica	Hadian et al. (2012) and Sefidkon and Jamzad (2005)
			S. parnassica subsp. parnassica	Michaelakis et al. (2007)
			S. pilosa subsp. origanita	Dardioti et al. (2012)
			S. pilosa subsp. pilosa	Konakchiev and Tsankova (2002)
			S. rechingeri	Hadian et al. (2012) and Sefidkon et al. (2007)
			S. sahendica	Gohari et al. (2011), Hassanpouraghdam et al. (2009), Sefidkon and Akbarinia (2009), Sefidkon et al. (2004), and Taherpour et al. (2008)
			S. spicigera	Eminagaoglu et al. (2007), Kotan et al. (2010), and Sefidkon and Jamzad (2004)
			S. spinosa	Michaelakis et al. (2007) and Skoula and Grayer (2005)
			S. subspicata	Skocibusic et al. (2006)
			S. thymbra	Economou et al. (2011), Figueredo et al. (2012), Giweli et al. (2012), Goren et al. (2004), Karpouhtsis et al. (1998), Loizzo et al. (2008b), Markovic et al. (2011), Michaelakis et al. (2007), Mullerriebau et al. (1995), Ozturk (2012), Piras et al. (2011), Schulz et al. (2005), Skoula and Grayer (2005), and Ugur et al. (2009)
			S. wiedemanniana	Altundag and Aslim (2011), Baser et al. (2001), Yucel and Aslim (2011)
			S. x delpozoi hybrid (S. intricata x S. obovata)	Jordan et al. (2010)
			p-Menth-3-en-8-ol	C10H18O	154.249298	S. viminea	Tucker et al. (2000) and Vila et al. (2000)
p-Mentha-3,8-diene	C10H16	136.233994	S. viminea	Vila et al. (2000)
Piperitenone	C10H14O	150.217606	S. dartvinii	Niemeyer (2010)
			S. fruticosa	Coelho et al. (2007, 2012)
			S. parvifolia	Cabana et al. (2013), Dambolena et al. (2009), Lima et al. (2011), and Luna et al. (2008)
			Piperitenone oxide	C10H14O2	166.216995	S. fruticosa	Coelho et al. (2007, 2012)
S. paradoxa	Tolossa et al. (2007)
S. parvifolia	Cabana et al. (2013), Lima et al. (2011), and Luna et al. (2008), Viturro et al. (2000)
S. paradoxa	Tolossa et al. (2007)
S. pseudosimensis	Vagionas et al. (2007)
Piperitone	C10H16O	152.233398				S. fruticosa	Coelho et al. (2012)
			S. parvifolia	Dambolena et al. (2009), Lima et al. (2011), Luna et al. (2008)
			Piperitone oxide	C10H16O2	168.232803	S. fruticosa	Coelho et al. (2012)
Pulegone	C10H16O	152.233398	S. abyssinica subsp. abyssinica	Tolossa et al. (2007)
			S. boliviana	Dambolena et al. (2009) and Hatami et al. (2011)
			S. brownei	Rojas and Usubillaga (2000)
			S. douglasii	Peer and Langenheim (1998)
			S. fruticosa	Coelho et al. (2007, 2012)
			S. odora	Fuselli et al. (2008)
			S. paradoxa	Tolossa et al. (2007)
			S. parvifolia	Cabana et al. (2013)
			S. viminea	Tucker et al. (2000) and Vila et al. (2000)
			rho-Cymene	C10H14	134.218201	S. intermedia	Sadeghi et al. (2013)
S. sahendica	Yousefzadi et al. (2012)
Sabinene	C10H16	136.233994				S. pseudosimensis	Tchoumbougnang et al. (2009)
			S. thymbra	Loizzo et al. (2008b)
			Spathulenol	C15H24O	220.350494	S. biflora	Vagionas et al. (2007)
S. cuneifolia	Skocibusic and Bezic (2004)
S. macrantha	Gohari et al. (2006a, 2006b)
S. masukensis	Vagionas et al. (2007)
S. parnassica subsp. parnassica	Tzakou and Skaltsa (2003)
S. pseudosimensis	Vagionas et al. (2007)
S. sahendica	Taherpour et al. (2008)
S. subspicata	Cavar et al. (2008) and Dunkic et al. (2012)
tau-Cadinol	C15H26O	222.366302				S. biflora	Matasyoh et al. (2007)
Terpinen-4-ol	C10H18O	154.249298	S. biflora	Vagionas et al. (2007)
			S. khuzestanica	Farsam et al. (2004)
			S. kitaibelii	Slavkovska et al. (2001)
			S. macrosiphonia	Amiri (2011)
			S. montana	Barbosa et al. (2012)
			S. montana subsp. montana	Slavkovska et al. (2001)
			S. montana subsp. pisidica	Slavkovska et al. (2001)
			S. thymbra	Dell'Agli et al. (2012)
			Terpineol	C10H18O	154.249298	S. macrosiphonia	Amiri (2011)
Tetradecane	C14H30	198.388	S. khuzestanica	Kheirandish et al. (2011)
Thymohydroquinone	C10H14O2	166.216995	S. hortensis	Taborsky et al. (2012)
			S. montana	Taborsky et al. (2012)
			S. thymbra	Fleisher and Fleisher (2005)
			Thymol	C10H14O	150.217606	S. aintabensis	Sampson et al. (2005)
S. atropatana	Gohari et al. (2005) and Moradi et al. (2008)
S. bachtiarica	Ahanjan et al. (2011a), Hadian et al. (2012), Hashemi et al. (2011), Mohammadpour et al. (2012), Pirbalouti et al. (2013), and Sefidkon and Jamzad (2000)
S. boliviana	Hatami et al. (2011)
S. cilicica	Schulz et al. (2005)
S. cuneifolia	Altun and Goren (2007), Biavati et al. (2004), Kan et al. (2006), Kosar et al. (2008), Oke et al. (2009a), Orhan et al. (2011), Schulz et al. (2005), and Tumen et al. (1998b)
S. edmondi	Sefidkon and Jamzad (2006)
S. hortensis	Adiguzel et al. (2007), Azaz et al. (2005), Baser et al. (2004), Deighton et al. (1994), Dikbas et al. (2010, 2011), Gulluce et al. (2003), Heydarzade and Moravvej (2012), Hoferl et al. (2009), Khajeh (2011), Kizil (2009), Kizil et al. (2009), Kubatova et al. (2001), Razzaghi-Abyaneh et al. (2008), Soran and Lung (2010), and Soran et al. (2011)
S. horvatii	Lakusic et al. (2008, 2011)
S. horvatii subsp. macrophylla	Dardioti et al. (2010)
S. intermedia	Sadeghi et al. (2013) and Sefidkon and Jamzad (2005)
S. khuzestanica	Farsam et al. (2004) and Saei-Dehkordi et al. (2012)
S. laxiflora	Sonboli et al. (2004)
S. macrantha	Sefidkon and Jamzad (2005)
S. mittica	Gohari et al. (2005)
S. montana	Barbosa et al. (2012), Bezbradica et al. (2005), Cavar et al. (2008); Coelho et al. (2012), Damjanovic-Vratnica et al. (2011), de Oliveira et al. (2011, 2012), Djenane et al. (2012), Dunkic et al. (2012), Fraternale et al. (2007), Grosso et al. (2009a, 2009b), Haloci et al. (2011), Ibraliu et al. (2011), Kulisic et al. (2005), Mastelic and Jerkovic (2003), Mirjana and Nada (2004), Palavra et al. (2011), Penalver et al. (2005), Prieto et al. (2007), Radonic and Milos (2003b), Serrano et al. (2011), Skocibusic and Bezic (2003, 2004, Stoilova et al. (2008), and Vidic et al. (2009)
S. montana subsp. montana	Nhu-Trang et al. (2006)
S. montana subsp. variegata	Dunkic et al. (2010)
S. mutica	Hadian et al. (2012) and Sefidkon and Jamzad (2005)
S. parnassica subsp. parnassica	Michaelakis et al. (2007)
S. pilosa subsp. origanita	Dardioti et al. (2012)
S. pilosa subsp. pilosa	Konakchiev and Tsankova (2002)
S. pseudosimensis	Vagionas et al. (2007)
S. robusta	Sefidkon and Akbarinia (2009) and Tchoumbougnang et al. (2009)
S. sahendica	Gohari et al. (2011), Sefidkon et al. (2004), Taherpour et al. (2008), and Yousefzadi et al. (2012)
S. spicigera	Eftekhar et al. (2009), Gohari et al. (2006a, 2006b, 2012), Kotan et al. (2010), and Sefidkon and Jamzad (2004)
S. spinosa	Michaelakis et al. (2007) and Skoula and Grayer (2005)
S. subspicata	Cavar et al. (2008)
S. subspicata subsp. liburnica	Dunkic et al. (2007)
S. subspicata subsp. subspicata	Dunkic et al. (2007)
S. thymbra	Dell'Agli et al. (2012), Figueredo et al. (2012), Fleisher and Fleisher (2005), Giweli et al. (2012), Goren et al. (2004), Karousou et al. (2005), Karpouhtsis et al. (1998), Lagouri et al. (1993), Loizzo et al. (2008b), Markovic et al. (2011), Michaelakis et al. (2007), Muller-Riebau et al. (1997), Mullerriebau et al. (1995), Ozturk (2012), Piras et al. (2011), and Skoula and Grayer (2005)
S. wiedemanniana	Baser et al. (2001) and Sampson et al. (2005)
S. x delpozoi hybrid (S. intricata x S. obovata)	Jordan et al. (2010)
Thymol methyl ether	C11H16O	164.244095				S. montana	Radonic and Milos (2003b) and Skocibusic and Bezic (2004)
			S. spicigera	Kotan et al. (2010)
			S. subspicata	Skocibusic et al. (2006)
			S. thymbra	Michaelakis et al. (2007)
			Thymoquinone  (p-cymene-2,5-dione)	C10H12O2	164.201096	S. atropatana	Moradi et al. (2008)
S. cuneifolia	Altun and Goren (2007)
S. fruticosa	Coelho et al. (2012)
S. hortensis	Taborsky et al. (2012)
S. montana	Coelho et al. (2012), Grosso et al. (2009b), Mastelic and Jerkovic (2003), Palavra et al. (2011), and Taborsky et al. (2012)
S. spicigera	Gohari et al. (2012)
S. thymbra	Fleisher and Fleisher (2005)
trans-Sabinene hydrate	C10H18O	154.249298				S. horvatii subsp. macrophylla	Dardioti et al. (2010)
			S. montana subsp. montana	Slavkovska et al. (2001)
			S. montana subsp. montana	Slavkovska et al. (2001)
			S. kitaibelii	Slavkovska et al. (2001)
			Viridiflorol	C15H26O	222.366302	S. visianii	Vidic et al. (2009)
α-Bisabolol	C15H26O	222.366302	S. punctata subsp. punctata	Tariku et al. (2010)
α-Bisabolol oxide-B	C15H26O2	238.365707	S. biflora	Vagionas et al. (2007)
α-Cadinol	C15H26O	222.366302	S. biflora	Matasyoh et al. (2007)
α-Copaene	C15H24	204.351105	S. subspicata	Bezic et al. (2009)
α-Eudesmol	C15H26O	222.366302	S. isophylla	Habibi et al. (2007), Sefidkon and Jamzad (2006)
			S. subspicata	Dunkic et al. (2012)
			α-Pinene	C10H16	136.233994	S. cuneifolia	Mirjana et al. (2004), Savikin et al. (2010), Skocibusic and Bezic (2004), Tommasi et al. (2008, 2009
S. hortensis	Kizil (2009); Pistelli et al. (2013), Soran et al. (2011)
S. kitaibelii	Slavkovska et al. (2001)
S. montana subsp. montana	Slavkovska et al. (2001)
S. montana subsp. pisidica	Slavkovska et al. (2001)
S. sahendica	Taherpour et al. (2008)
S. subspicata	Bezic et al. (2009) and Skocibusic et al. (2006)
S. subspicata subsp. liburnica	Dunkic et al. (2007)
S. subspicata subsp. subspicata	Dunkic et al. (2007)
S. thymbra	Loizzo et al. (2008b)
S. visianii	Bezic et al. (2009)
α-Terpinene	C10H16	136.233994				S. cuneifolia	Altun and Goren (2007)
			S. edmondi	Masumeh (2006)
			S. hortensis	Alizadeh et al. (2010a), Dikbas et al. (2011), Pfefferkorn et al. (2008), Pistelli et al. (2013), Rezvanpanah et al. (2011), Sajfrtova et al. (2013), and Tozlu et al. (2011)
			S. horvatii	Lakusic et al. (2008)
			S. montana subsp. montana	Slavkovska et al. (2001)
			S. sahendica	Hassanpouraghdam et al. (2009)
			S. subspicata	Bezic et al. (2009)
			S. thymbra	Hassanpouraghdam et al. (2009)
			α-Terpineol	C10H18O	154.249298	S. edmondi	Sefidkon and Jamzad (2006)
S. hortensis	Kizil (2009)
S. montana	Lampronti et al. (2006) and Mastelic and Jerkovic (2003)
S. pseudosimensis	Tchoumbougnang et al. (2009)
S. x delpozoi hybrid  (S. intricata x S. obovata)	Jordan et al. (2010)
α-Thujene	C10H16	136.233994				S. hortensis	Pistelli et al. (2013)
			S. horvatii	Lakusic et al. (2008)
			S. sahendica	Hassanpouraghdam et al. (2009)
			S. visianii	Bezic et al. (2009) and Dunkic and Bezic (2008)
			α-Thujone	C10H16O	152.233398	S. pomifera	Glamoclija et al. (2006)
S. rechingeri	Sefidkon et al. (2007)
β-Bisabolene	C15H24	204.351105				S. bachtiarica	Meshkatalsadat and Shabaninejad (2011)
β-Bourbonene	C15H24	204.351105	S. biflora	Matasyoh et al. (2007) and Vagionas et al. (2007)
			S. isophylla	Habibi et al. (2007)
			S. masukensis	Vagionas et al. (2007)
			β-Caryophyllene	C15H24	204.351105	S. bachtiarica	Sefidkon and Jamzad (2000)
S. biflora	Matasyoh et al. (2007)
S. boissieri	Sajjadi and Baluchi (2002)
S. boliviana	Hatami et al. (2011) and Viturro et al. (2000)
S. corulea	Azaz et al. (2002)
S. cuneifolia	Skocibusic and Bezic (2004)
S. hortensis	Pistelli et al. (2013), Tozlu et al. (2011)
S. horvatii	Lakusic et al. (2008)
S. isophylla	Habibi et al. (2007) and Sefidkon and Jamzad (2006)
S. montana	Barbosa et al. (2012), Fraternale et al. (2007), and Stoilova et al. (2008)
S. parnassica subsp. parnassica	Michaelakis et al. (2007)
S. soinosa	Michaelakis et al. (2007)
S. subspicata	Skocibusic et al. (2006)
S. thymbra	Economou et al. (2011), Fleisher and Fleisher (2005), Michaelakis et al. (2007), and Piras et al. (2011)
β-Cubebene	C15H24	204.351105				S. cuneifolia	Skocibusic and Bezic (2004)
β-Eudesmol	C15H26O	222.366302	S. isophylla	Sefidkon and Jamzad (2006)
			S. subspicata	Dunkic et al. (2012)
			β-Ocimene	C10H16	136.233994	S. biflora	Matasyoh et al. (2007)
β-Phellandrene	C10H16	136.233994	S. montana	Fraternale et al. (2007)
β-Pinene	C10H16	136.233994	S. biflora	Matasyoh et al. (2007)
β-Thujone	C10H16O	152.233398	S. pomifera	Glamoclija et al. (2006)
γ-Eudesmol	C15H26O	222.366302	S. isophylla	Habibi et al. (2007), Sefidkon and Jamzad (2006)
γ-Terpinene	C10H16	136.233994	S. bachtiarica	Pirbalouti et al. (2013), Sefidkon and Jamzad (2000)
			S. boissieri	Kurkcuoglu et al. (2001) and Sajjadi and Baluchi (2002)
			S. boliviana	Viturro et al. (2000)
			S. cilicica	Schulz et al. (2005)
			S. cuneifolia	Altun and Goren (2007), Bezic et al. (2009), Biavati et al. (2004), Kan et al. (2006), Oke et al. (2009a), Orhan et al. (2011), and Schulz et al. (2005)
			S. edmondi	Masumeh (2006) and Sefidkon and Jamzad (2006)
			S. hortensis	Adiguzel et al. (2007), Alizadeh et al. (2010a), Alizadeh-Salteh et al. (2010b), Annan et al. (2013), Baher et al. (2002), Dawidowicz and Rado (2010), Dikbas et al. (2011), Figueredo et al. (2012), Ghannadi (2002), Gulluce et al. (2003), Hadian et al. (2010a), Heydarzade and Moravvej (2012), Khajeh (2011), Kizil et al. (2009), Mihajilov-Krstev et al. (2009b), Nedorostova et al. (2011), Novak et al. (2006), Pavela (2009), Pfefferkorn et al. (2008), Pistelli et al. (2013), Rezvanpanah et al. (2011), Sajfrtova et al., 2013, Sampson et al. (2005), Sefidkon et al. (2006), Shatar and Altantsetseg (2000), Tozlu et al. (2011), and Viturro et al. (1999)
			S. horvatii	Lakusic et al. (2008)
			S. intermedia	Sadeghi et al. (2013), Sefidkon and Jamzad (2005)
			S. khuzestanica	Farsam et al. (2004), Hadian et al. (2012), Sefidkon and Ahmadi (2000), and Sefidkon and Jamzad (2005)
			S. kitaibelii	Slavkovska et al. (2001)
			S. laxiflora	Sonboli et al. (2004)
			S. montana	Barbosa et al. (2010, 2012, Bezic et al. (2009), Chizzola (2003), Coelho et al. (2012), Damjanovic-Vratnica et al. (2011), Djenane et al. (2012), Grosso et al. (2009a), Ibraliu et al. (2011), Marin et al. (2012), Mastelic and Jerkovic (2003), Michaelakis et al. (2007), Miladi et al. (2013), Mirjana and Nada (2004), Prieto et al. (2007), Radonic and Milos (2003b), Sfeir et al. (2013), Skocibusic and Bezic (2004), and Stoilova et al. (2008)
			S. montana subsp. montana	Nhu-Trang et al. (2006) and Slavkovska et al. (2001)
			S. montana subsp. pisidica	Slavkovska et al. (2001)
			S. mutica	Sefidkon and Jamzad (2005)
			S. parnassica subsp. parnassica	Michaelakis et al. (2007)
			S. sahendica	Gohari et al. (2011), Hassanpouraghdam et al. (2009), Sefidkon and Akbarinia (2009), Sefidkon et al. (2004), Taherpour et al. (2008), and Yousefzadi et al. (2012)
			S. spicigera	Eminagaoglu et al. (2007), Kotan et al. (2010), and Sefidkon and Jamzad (2004)
			S. spinosa	Skoula and Grayer (2005)
			S. subspicata	Skocibusic et al. (2006)
			S. thymbra	Cetin et al. (2010), Economou et al. (2011), Fleisher and Fleisher (2005), Giweli et al. (2012), Glamoclija et al. (2006), Goren et al. (2004), Karpouhtsis et al. (1998), Loizzo et al. (2008b), Markovic et al. (2011), Michaelakis et al. (2007), Mullerriebau et al. (1995), Ozturk (2012), Piras et al. (2011), Sampson et al. (2005), Schulz et al. (2005), Skoula and Grayer (2005), Ugur et al. (2009)
			S. wiedemanniana	Altundag and Aslim (2011), and Baser et al. (2001)
			S. x delpozoi hybrid (S. intricata x S. obovata)	Jordan et al. (2010)
			δ-Cadinene	C15H24	204.351105	S. biflora	Matasyoh et al. (2007)

As can be seen from Table 2, thymol and carvacrol are determined as main compounds for the majority of species. Numerous studies in the literature indicated that both compounds are especially the main volatiles for S. cuneifolia, S. hortensis, S. montana, S. spicigera, S. thymbra, and S. bachtiarica. Additionally, carvacrol is an important main compound for S. khuzestanica. Thymol and carvacrol are followed by p-cymene and γ-terpinene, respectively. According to several reports, both compounds are the main volatiles of S. cuneifolia, S. hortensis, S. montana, S. sahendica, and S. thymbra (see Table 3 for the references).

Table 3. Factors affecting essential oil composition of Satureja species and chemotypes suggested by researchers.

Effect of developmental stages on essential oil composition
Plant	Developmental stage/short explanation	Reference
S. hortensis	Floral budding, full flowering, immature fruit, and ripened fruit stages γ-Terpinene was the major compound of the oil at all developmental stages, except the ripened fruit stage where it was replaced by carvacrol (46.4%)	Saharkhiz et al. (2011)
S. horvatii	Before flowering, flowering, full stage of fruiting – Before flowering: Linalool (37.4%), thymol (27.3%), carvacrol (12.2%) – Flowering: Linalool (56.6–57.5%), thymol (15.5–15.8%), carvacrol (1.4–1.5%) – Full stage of fruiting: Linalool (58.4–65.8%), thymol (7.6–1.3%), carvacrol (0.7–0.1%)	Lakusic et al. (2011)
S. montana	Before flowering, after flowering stages Before flowering: Thymol (37.36%), carvacrol (15.47%), γ-terpinene (11.75%), p-cymene (7.86%) After flowering: Thymol (27.68%), carvacrol (4.40%), γ-terpinene (8.66%), p-cymene (31.37%)	Damjanovic-Vratnica et al. (2011)
	Before flowering, during flowering, after flowering stages – Before flowering: Carvacrol (52.4%), thymol (11.0%), γ-terpinene (5.8%), methyl carvacrol (4.6%), methyl thymol (3.2%), borneol (3.2%) – During flowering: p-Cymene (15.2%), carvacrol (26.2%), geraniol (10.2%), geranyl acetate (5.2%), borneol (6.5%), methyl carvacrol (6.4%), methyl thymol (3.3%), γ-terpinene (4.9%), thymol (5.4%) – After flowering: p-Cymene (25.6%), carvacrol (16.1%), methyl thymol (12.8%), methyl carvacrol (5.4%), borneol (11.5%), thymol (2.6%), γ-terpinene (-)	Mirjana and Nada (2004)
	S. macrosiphonia	Pre-flowering stage, flowering stages – Pre-flowering stage: Terpinen-4-ol (20.8%), terpineol (14.7%). – Flowering stage: Terpineol (26.7%), borneol (16.6%).	Amiri (2011)
S. sahendica	Before flowering, full flowering, end of flowering stages – Before flowering: γ-Terpinene (42.3%), thymol (34.6%), p-cymene (11.7%). – Full flowering: p-Cymene (30.2%), thymol (29.6), γ-terpinene (27.7%). – End of flowering: p-Cymene (58.1%), thymol (19.3%), γ-terpinene (9.3%).	Sefidkon and Akbarinia (2009)
S. cuneifolia	Preflowering, flowering, post flowering stages Thymol (42.5–45.2%), p-cymene (19.4–24.3%), carvacrol (8.5–13.2%) are the main compounds at all stages	Kosar et al. (2008)
	Before flowering, during flowering, after flowering stages Linalool and borneol appeared to be relatively constant but carvacrol, limonene and a-pinene showed variability during ontogenetic stages.	Mirjana et al. (2004)
	S. thymbra	Just before flowering, full flowering – Just before flowering: p-Cymene (10.39%), γ-terpinene (20.12%), thymol (42.15%), thymol methyl ether (4.12%) – Full flowering: p-Cymene (9.19%), γ-terpinene (14.64%), thymol (24.32%), carvacrol (30.39%), β-caryophyllene (4.89%)	Michaelakis et al. (2007)
February, March, April, May, June and July – February: p-Cymene (95.4 %, v/w), γ-terpinene (21.5 %, v/w), 1,8-cineole (14.7 %, v/w) – March: p-Cymene (77.4 %, v/w), γ-terpinene (53.8 %, v/w), 1,8-cineole (20.4 %, v/w) – April: p-Cymene (31.8 %, v/w), γ-terpinene (168.4 %, v/w), 1,8-cineole (9.9 %, v/w) – May: p-Cymene (30.4 %, v/w), γ-terpinene (103.7 %, v/w), 1,8-cineole (11.3 %, v/w) – June: p-Cymene (58.5 %, v/w), γ-terpinene (128.5 %, v/w), 1,8-cineole (9.3 %, v/w) – July: p-Cymene (74.1 %, v/w), γ-terpinene (139.2 %, v/w), 1,8-cineole (5.0 %, v/w)		Muller-Riebau et al. (1997)
S. rechingeri		Beginning of flowering, full flowering – Beginning of flowering: Carvacrol (56.1%), p-cymene (14.0%), α-thujone (4.7%). – Full flowering: Carvacrol (84.0–89.3%).	Sefidkon et al. (2007)
S. spinosa	Before flowering, during flowering – Before flowering: linalool (61.5%) – During flowering: linalool (47.4%)	Aslan et al. (2004)
S. parnassica subsp. parnassica	Flowering and vegetative stages In both stages spathulenol, p-cymene and linalool were main compounds	Tzakou and Skaltsa (2003)
Effect of isolation techniques on essential oil composition
Plant	Techniques used/ short explanation	Reference
S. hortensis	Microwave extraction, hydrodistillation – Microwave extraction: p-Cymene (10.95%), γ-terpinene (8.33%), carvacrol (59.35%) – Hydrodistillation: p-Cymene (19.93%), γ-terpinene (20.53%), carvacrol (38.83%)	Figueredo et al. (2012)
	Hydrodistillation, Microwave Assisted Hydrodistillation – Hydrodistillation: Carvacrol (48.69%), γ-terpinene (31.95%), α-terpinene (3.89%) – Microwave Assisted Hydrodistillation: Carvacrol (47.74%), γ-terpinene (29.99%), α-terpinene (4.55%)	Rezvanpanah et al. (2011)
	Maceration, sonication, microwave-assisted extraction Best results were obtained by maceration with methanol.	Soran and Lung (2010)
	Steam distillation, pressurized liquid extraction, matrix solid-phase dispersion – Steam distillation: Carvacrol (69.2%), γ-terpinene (22.0%), p-cymene (3.9%) – Pressurized liquid extraction: Carvacrol (66.7%), γ-terpinene (23.4%), p-cymene (3.7%) – Matrix solid-phase dispersion: Carvacrol (71.5%), γ-terpinene (19.4%), p-cymene (2.9%)	Dawidowicz and Rado (2010)
	Hydro distillation, water and steam distillation, direct steam distillation – Hydro distillation: Carvacrol (46.0%), γ-terpinene (37.7%) – Water and steam distillation: Carvacrol (44.0%), γ-terpinene (41.8%) – Direct steam distillation: Carvacrol (12.3%), γ-terpinene (70.4%)	Sefidkon et al. (2006)
	S. ftuticosa	Hydrodistillation, supercritical fluid extraction – Hydrodistillation: Pulegone (40.6%), isomenthone (20.7%), piperitone oxide + piperitone (10.5%), piperitenone (9.0%), Piperitenone oxide (6.8%) – Supercritical fluid extraction: Pulegone (35.7%), isomenthone (17.6%), piperitone oxide + piperitone (9.8%), piperitenone (11.2%), Piperitenone oxide (10.7%)	Coelho et al. (2012)
Hydrodistillation, supercritical fluid extraction – Hydrodistillation: Pulegone (40.0–41.0%), isomenthone (21.0%), piperitone (9.0–11.0%), piperitone oxide (7.0–9.0%) – Supercritical fluid extraction: Pulegone (33.0–36.0%), isomenthone (14.0–18.0%), piperitone (11.0–12.0%), piperitone oxide (11.0–13.0%)		Coelho et al. (2007)
S. montana		Hydrodistillation, supercritical fluid extraction – Hydrodistillation: p-Cymene (12.8%), γ-terpinene (8.9%), thymol (11.0%), carvacrol (52.2%) – Supercritical fluid extraction: p-Cymene (10.1%), γ-terpinene (4.3%), thymol (10.9%), carvacrol (52.7%)	Coelho et al. (2012)
	Hydrodistillation, supercritical fluid extraction – Hydrodistillation: Carvacrol (52.2–62.0%), thymol (8.6–11.0%), p-Cymene (6.9–12.8%), γ-terpinene (6.4–9.4%), p-bisabolene (2.0–2.7%), thymoquinone (0.2%) – Supercritical fluid extraction: Carvacrol (41.7–64.5%), thymol (6.0–11.3%), p-Cymene (6.0–17.8%), γ-terpinene (2.3–6.0%), p-bisabolene (2.2–3.5%), thymoquinone (1.6–3.0%)	Grosso et al. (2009a)
	S. thymbra	Microwave extraction, hydrodistillation – Microwave extraction: Carvacrol (38.72%), thymol (17.15%), p-cymene (12.26%) – Hydrodistillation: Carvacrol (34.03%), thymol (14.36%), p-cymene (18.32%S)	Figueredo et al. (2012)
Hydrodistillation, thermal desorption, headspace GC-MS All showed similar results		Goren et al. (2004)
S. sahendica		Hydrodistillation, steam distillation – Hydrodistillation: Thymol (37.2%), p-cymene (32.6%), γ-terpinene (11.5%). – Steam distillation: Thymol (66.0%), p-cymene (20.3%), γ-terpinene (3.6%).	Gohari et al. (2011)
Effect of climatic conditions on essential oil composition
Plant	Climatic conditions/short explanation	Reference
S pilosa subsp. origanita	Meso-Mediterranean and Temperate Axeric bioclimates The essential oil content and the amount of carvacrol gradually decrease from the Meso-Mediterranean to the Temperate Axeric bioclimates, while the amounts of p-cymene, thymol and linalool increase	Dardioti et al. (2012)
S. hortensis	Summer and autumn Summer: carvacrol (55.3–59.5), γ-terpinene (30.1–33.8) Autumn: carvacrol (55.5–57.9), γ-terpinene (32.1–33.5)	Pfefferkorn et al. (2008)
S. horvatii subsp. macrophylla	Mediterranean, SubMediterranean and Temperate Axeric bioclimates – Carvacrol dominates in areas with Mediterranean and SubMediterranean bioclimates – Linalool or trans-sabinene hydrate and/or borneol in SubMediterranean and Temperate Axeric bioclimates – Thymol in Mediterranean, SubMediterranean and Temperate Axeric bioclimates – The essential oil negatively related to the altitude	Dardioti et al. (2010)
Effect of drying methods on essential oil composition
Plant	Drying methods/short explanation	Reference
S. hortensis	Drying by microwave at two temperatures: 50 and 70 °C, and six microwave powers: 100, 180, 300, 450, 600, and 900 W Maximum carvacrol content (63.9%) obtained at 300 W microwave drying. Maximum γ-terpinene content (28.2%) obtained at drying by 70 °C that it had little difference with 50 °C, 100 and 300 W.	Ebadi et al. (2011)
	Sun drying, shade drying and oven drying Sun drying: carvacrol (46.8), γ-terpinene (39.4) Shade drying: carvacrol (46.0), γ-terpinene (37.7) Oven drying: carvacrol (48.1), γ-terpinene (38.4)	Sefidkon et al. (2006)
	Differences between wild and cultivated plants in terms of essential oil composition
Plant	Short explanation	Reference
S. hortensis	In vitro plant: p-cymene (15.6%), γ-terpinene (29%), carvacrol (14.4%), caryophyllene (20.0%), β-bisabolene (4.0%), α-pinene (1.5%), α-thujene (1.5%), α-terpinene (2.5%)\ Field grown plant: p-cymene (30.3%), γ-terpinene (11.7%), carvacrol (-), caryophyllene (5.0%), β-bisabolene (3.5%), α-pinene (3.0%), α-thujene (1.4%), α-terpinene (3.4%)	Pistelli et al. (2013)
S. khuzestanica	Wild: carvacrol (93.9), eugenol (1.0), p-cymene (0.8), thymol (0.6) Cultivated: carvacrol (80.6), p-cymene (4.8), myrcene (1.5), γ-terpinene (2.1), terpinene-4-ol (2.1)	Farsam et al. (2004)
Chemotypes suggested by researchers
Plant	Short explanation	Reference
S. montana	Carvacrol and thymol chemotypes were suggested	Dunkic et al. (2012)
	Linalool and carvacrol/p-cymene chemotypes were suggested	Chizzola (2003)
	S. subspicata	α-Eudesmol, β-eudesmol and spathulenol chemotypes were suggested	Dunkic et al. (2012)
S pilosa subsp. origanita	Oils rich in carvacrol (oregano odor) (up to 62.3), oils rich in p-cymene (thyme odor) (up to 49.8), oils rich in thymol (thyme odor) (up to 48.1), oils rich in linalool (levander odor) (up to 59.2 and 82.7) were suggested	Dardioti et al. (2012)
S. cuneifolia	Linalool, borneol, α-pinene and borneol/α-pinene chemotypes were suggested	Tommasi et al. (2008)
S. biflora	Linalool chemotype was suggested	Matasyoh et al. (2007)
S. subspicata subsp. subspicata	α-Pinene chemotype was suggested	Dunkic et al. (2007)
S. subspicata subsp. liburnica	α-Pinene chemotype was suggested	Dunkic et al. (2007)
S. thymbra	Thymol, carvacrol chemotypes were suggested	Fleisher and Fleisher (2005)
S. douglasii	Bicyclic or isomenthone chemotypes were suggested	Peer and Langenheim (1998)

According to the reports presented in Table 3, some Satureja species have unusual chemical profiles. For example; piperitone, piperitenone, and piperitenone oxide are the main compounds of S. parvifolia and S. fruticosa Beguinot (Cabana et al., 2013; Coelho et al., 2007, 2012; Dambolena et al., 2009; Lima et al., 2011; Luna et al., 2008; Viturro et al., 2000). Additionally, camphor was determined as the major compound of S. visianii Silic (Bezic et al., 2009; Dunkic & Bezic, 2008; Vidic et al., 2009). It is possible to increase the number of these kinds of examples as the following: isomenthone for S. boliviana (Dambolena et al., 2009; Hatami et al., 2011; Senatore et al., 1998), thymoquinone (Grosso et al., 2009b; Mastelic & Jerkovic, 2003; Palavra et al., 2011; Taborsky et al., 2012), geraniol (Cavar et al., 2008; Mastelic & Jerkovic, 2003; Mirjana & Nada, 2004; Radonic & Milos, 2003b), and carvacrol methyl ether (Serrano et al., 2011; Skocibusic & Bezic, 2004) for S. montana.

In addition to the compounds discussed above, linalool, borneol, β-caryophyllene, α-pinene, limonene, pulegone, α-terpinene, terpinen-4-ol, spathulenol, and myrcene are reported as the major compounds of Satureja species. These findings are supported by many reports presented in Table 3.

As it is well known, essential oil composition of plant species is strictly influenced by environmental factors such as climatic conditions, soil type, altitude, and temperature. Any assessments made by ignoring these factors may be misleading. Therefore, factors affecting essential oil composition of Satureja species and chemotypes suggested by researchers are presented in Table 3. In the table, the effect of developmental stages on essential oil composition, isolation techniques, climatic conditions, and drying methods is presented without any discussion. Additionally, in terms of essential oil compositions and chemotype, differences between wild and cultivated plants suggested by researchers are presented in table.

Phenolic acids

Phenolic acids determined in Satureja species are presented in Table 4 together with their molecular formula and average mass (Da) values. Characteristic phenolic acids found is Satureja species are also presented in Figure 2.

Figure 2. Characteristic phenolic acids of Satureja species.

Table 4. Phenolic acids of Satureja species.

Major Terpenoids	Molecular formula	Average mass (Da)	Plant Species	Reference
Rosmarinic acid	C18H16O8	360.314789	S. aintabensis	Askun et al. (2013)
			S. atropatana	Shekarchi et al. (2012) and Hajimehdipoor et al. (2010)
			S. bachtiarica	Shekarchi et al. (2012), Hajimehdipoor et al. (2010), and Ebrahimi et al. (2008)
			S. cilicica	Cetojevic-Simin et al. (2012)
			S. corulea	Cetojevic-Simin et al. (2012)
			S. cuneifolia	Kosar et al. (2003)
			S. edmondi	Ebrahimi et al. (2008)
			S. hortensis	Ishigaki et al. (2013), Lung et al. (2013), Plander et al. (2012), Shekarchi et al. (2012), Hajimehdipoor et al. (2010), Hadian et al. (2010a), Bros et al. (2009), Tepe and Sokmen (2007), Kosar et al. (2005), Exarchou et al. (2002), and Bertelsen et al. (1995)
			S. icerica	Cetojevic-Simin et al. (2012)
			S. khuzestanica	Shekarchi et al. (2012), Hadian et al. (2011); Hajimehdipoor et al. (2010), and Ebrahimi et al. (2008)
			S. kitaibelii	Stanojkovic et al. (2013)
			S. laxiflora	Ebrahimi et al. (2008)
			S. macrantha	Shekarchi et al. (2012) and Hajimehdipoor et al. (2010)
			S. montana	da Silva et al. (2013)
			S. mutica	Shekarchi et al. (2012), Hajimehdipoor et al. (2010), and Ebrahimi et al. (2008)
			S. parvifolia	Cabana et al. (2013)
			S. rechingeri	Ebrahimi et al. (2008)
			S. sahendica	Ebrahimi et al. (2008)
			S. spicigera	Ebrahimi et al. (2008)
			S. subspicata	Stefan et al. (2014)
			p-Coumaric acid	C9H8O3	164.158005	S. aintabensis	Askun et al. (2013)
S. cilicica	Cetojevic-Simin et al. (2012)
S. corulea	Cetojevic-Simin et al. (2012)
S. icerica	Cetojevic-Simin et al. (2012)
Ferulic acid	C10H10O4	194.184006				S. aintabensis	Askun et al. (2013)
			S. cilicica	Cetojevic-Simin et al. (2012)
			S. corulea	Cetojevic-Simin et al. (2012)
			S. icerica	Cetojevic-Simin et al. (2012)
			Caffeic acid	C9H8O4	180.157394	S. cilicica	Cetojevic-Simin et al. (2012)
S. corulea	Cetojevic-Simin et al. (2012)
S. hortensis	Lung et al. (2013), Plander et al. (2012), and Kosar et al. (2005)
S. icerica	Cetojevic-Simin et al. (2012)
S. montana	Giao et al. (2009)
Gallic acid	C7H6O5	170.119507				S. cilicica	Cetojevic-Simin et al. (2012)
			S. corulea	Cetojevic-Simin et al. (2012)
			S. icerica	Cetojevic-Simin et al. (2012)
			Chlorogenic acid	C16H18O9	354.308685	S. montana	Palavra et al. (2011), Silva et al. (2009), and Giao et al. (2009)
S. subspicata	Stefan et al. (2014)
Protocatechuic acid	C7H6O4	154.120102				S. montana	Palavra et al. (2011) and Silva et al. (2009)
Vanillic acid	C8H8O4	168.146698	S. montana	Palavra et al. (2011) and Silva et al. (2009)
Carnosic acid	C20H28O4	332.433899	S. cuneifolia	Kosar et al. (2003)

As can be seen from Table 4, the most common phenolic acid found in Satureja species is rosmarinic acid. Reports in this section are particularly focused on S. hortensis, S. khuzestanica, S. mutica Fisch. & C. A. Mey., and S. bachtiarica. These are followed by S. atropatana and S. macrantha (C. A. Mey).

Caffeic acid is another phytochemical found in Satureja species. It was mainly found as one of the characteristic phenolic acids of S. cilicica P. H. Davis, S. hortensis, S. icerica L., and S. montana (Cetojevic-Simin et al., 2012; Giao et al., 2009; Kosar et al., 2005; Lung et al., 2013).

Other phenolic acids determined in these species are p-coumaric, ferulic, gallic, chlorogenic, protocatechuic, vanillic, and carnosic acids.

Flavonoids and related compounds

Flavonoids and related compounds found in Satureja species are presented in Table 5. The most common flavonoids are also given as Figure 3.

Figure 3. Characteristic flavonoids of Satureja species.

Table 5. Flavonoids and related compounds of Satureja species.

Compound	Molecular formula	Average mass (Da)	Plant Species	Reference
Flavones
Luteolin	C15H10O6	286.236298	S. aintabensis	Askun et al. (2013)
			S. atropatana	Gohari et al. (2009) and Moradi et al. (2008)
			S. cilicica	Cetojevic-Simin et al. (2012)
			S. corulea	Cetojevic-Simin et al. (2012)
			S. hortensis	Lung et al. (2013), Bros et al. (2009), and Exarchou et al. (2002)
			S. icerica	Cetojevic-Simin et al. (2012)
			S. obovata var. obovata	de Rojas et al. (1996a)
			S. obovata subsp. obovata var. obovata	de Rojas et al. (1996b)
			S. sahendica	Sharifi et al. (2011)
			S. parvifolia	van Baren et al. (2006)
			Apigenin	C15H10O5	270.236908	S. aintabensis	Askun et al. (2013)
S. cilicica	Cetojevic-Simin et al. (2012)
S. corulea	Cetojevic-Simin et al. (2012)
S. hortensis	Lung et al. (2013), Kosar et al. (2005), and Exarchou et al. (2002)
S. icerica	Cetojevic-Simin et al. (2012)
S. khuzestanica	Malmir et al. (2012)
S. sahendica	Hadjmohammadi et al. (2012) and Sharifi et al. (2011)
S. spinosa	Skoula et al. (2005)
Cirsilineol	C18H16O7	344.315399				S. khuzestanica	Malmir et al. (2012)
			S. spinosa	Skoula et al. (2005)
			Cirsimaritin	C17H14O6	314.28949	S. khuzestanica	Malmir et al. (2012)
S. spinosa	Skoula et al. (2005)
Genkwanin	C16H14O5	286.279388				S. thymbra	Skoula et al. (2005)
			S. spinosa	Skoula et al. (2005)
			Xanthomicrol	C18H16O7	344.315399	S. khuzestanica	Malmir et al. (2012)
S. spinosa	Skoula et al. (2005)
6-Hydroxyluteolin 7,3′-dimethyl ether	C17H14O7	330.28891				S. khuzestanica	Malmir et al. (2012)
			S. thymbra	Skoula et al. (2005)
			S. spinosa	Skoula et al. (2005)
			4′,5,6-Trihydroxy-3′,7-dimethoxyflavone	C17H14O7	330.28891	S. khuzestanica	Moghaddam et al. (2007)
5,6-Dihydroxy-7,8,3′,4′-tetramethoxyflavone or 5-desmethoxynobiletin (5-desmethoxynobiletine)	C19H18O8	374.34	S. mexicana	Estrada-Reyes et al. (2008)
			S. atropatana	Gohari et al. (2009) and Moradi et al. (2008)
			5-Hydroxy-7,4′-dimethoxyflavone	C17H14O5	298.2901	S. macrostema	Gutierrez (2013)
5,6,3′-Trihydroxy-7,8,4′-trimethoxyflavone	C18H16O8	360.314789	S. atropatana	Gohari et al. (2009) and Moradi et al. (2008)
6-Hydroxyluteolin	C15H10O7	302.235687	S. khuzestanica	Malmir et al. (2012)
6-Hydroxyluteolin 7,3′,4′-trimethyl ether	Not found	Not found	S. thymbra	Skoula et al. (2005)
8-Methoxycirsilineol	C19H18O8	374.3414	S. spinosa	Skoula et al. (2005)
Acacetin	C16H12O5	284.263489	S. khuzestanica	Malmir et al. (2012)
Diosmetin	C16H12O6	300.262909	S. khuzestanica	Malmir et al. (2012)
Diosmin	C28H32O15	608.544983	S. macrostema	Gutierrez (2013)
Ladanein	C17H14O6	314.28949	S. thymbra	Skoula et al. (2005)
Luteolin 7-methyl ether	C16H12O6	300.262909	S. thymbra	Skoula et al. (2005)
Thymonin (5,6,4′-trihydroxy-7,8,3′-trimethoxyflavone)	C18H16O8	360.314789	S. atropatana	Gohari et al. (2009) and Moradi et al. (2008)
			S. spinosa	Skoula et al. (2005)
			Thymusin	Not found	Not found	S. spinosa	Skoula et al. (2005)
Flavonols
Quercetin	C15H10O7	302.235687	S. aintabensis	Askun et al. (2013)
			S. cilicica	Cetojevic-Simin et al. (2012)
			S. corulea	Cetojevic-Simin et al. (2012)
			S. hortensis	Exarchou et al. (2002)
			S. icerica	Cetojevic-Simin et al. (2012)
			S. montana	da Silva et al. (2013)
			S. sahendica	Sharifi et al. (2011)
			Epicatechin-3-O-gallate	C22H18O10	442.372314	S. parvifolia	Cabana et al. (2013)
Epigallocatechin-3-O-gallate	C22H18O11	458.371704	S. parvifolia	Cabana et al. (2013)
Kaempferol	C15H10O6	286.236298	S. sahendica	Sharifi et al. (2011)
Myrcetin	C15H10O8	318.235107	S. sahendica	Sharifi et al. (2011)
Flavanones
Naringenin	C15H12O5	272.252808	S. aintabensis	Askun et al. (2013)
			S. cilicica	Cetojevic-Simin et al. (2012)
			S. corulea	Cetojevic-Simin et al. (2012)
			S. hortensis	Lung et al. (2013)
			S. icerica	Cetojevic-Simin et al. (2012)
			S. khuzestanica	Malmir et al. (2012)
			S. mexicana	Estrada-Reyes et al. (2008)
			S. obovata var. obovata	de Rojas et al. (1996a)
			S. obovata subsp. obovata var. obovata	de Rojas et al. (1996b)
			S. thymbra	Skoula et al. (2005)
			S. spinosa	Skoula et al. (2005)
			Eriodictyol (5,7,3′,4′-tetrahydroxyflavanone)	C15H12O6	288.252197	S. cilicica	Cetojevic-Simin et al. (2012)
S. corulea	Cetojevic-Simin et al. (2012)
S. hortensis	Kosar et al. (2005)
S. icerica	Cetojevic-Simin et al. (2012)
S. obovata var. obovata	de Rojas et al. (1996a)
S. obovata subsp. obovata var. obovata	de Rojas et al. (1996b)
S. obovata	de Rojas et al. (1999)
S. parvifolia	van Baren et al. (2006)
S. spinosa	Skoula et al. (2005)
S. thymbra	Skoula et al. (2005)
5,7,3′,5'-Tetrahydroxy flavanone	C15H12O6	288.252197				S. spicigera	Gohari et al. (2012)
5,4′-Dihydroxy-3′-methoxyflavanone-7-(6″-O-alpha-L-rhamnopyranosyl)-Beta-D-glucopyranoside	C28H34O14	594.561218	S. spicigera	Gohari et al. (2012)
Flavanone trimethyl ether	Not found	Not found	S. spinosa	Skoula et al. (2005)
Flavan-3-ols
Catechin	C15H14O6	290.268097	S. cilicica	Cetojevic-Simin et al. (2012)
			S. corulea	Cetojevic-Simin et al. (2012)
			S. icerica	Cetojevic-Simin et al. (2012)
			S. montana	Palavra et al. (2011) and Silva et al. (2009), Cetkovic et al. (2007b)
			Epicatechin	C15H14O6	290.268097	S. cilicica	Cetojevic-Simin et al. (2012)
S. corulea	Cetojevic-Simin et al. (2012)
S. icerica	Cetojevic-Simin et al. (2012)
S. montana	Cetkovic et al. (2007b)
Flavanonols
Aromadendrin	C15H12O6	288.252197	S. khuzestanica	Malmir et al. (2012)
			S. spinosa	Skoula et al. (2005)
			S. thymbra	Skoula et al. (2005)
			Taxifolin	C15H12O7	304.251587	S. khuzestanica	Malmir et al. (2012)
S. spinosa	Skoula et al. (2005)
S. thymbra	Skoula et al. (2005)
Flavanoid glucosides
5-Hydroxy-3,6,4′-trimethoxyflavonol-7-C-α-L-rhamnopyranosyl-(1 → 3)-β-D-glucopyranoside	Not found	Not found	S. macrostema	Gutierrez (2013)
4′-Methoxy-5,7,3′,5′-tetra-hydroxyflavanone-3-O-β-D-rhamnopyranosyl-(1 → 2)-β-d-rhamnopyranoside	Not found	Not found	S. macrostema	Gutierrez (2013)
5,4′-Dimethoxy-7,3′,5′-trihydroxyflavanone-3-O-β-d-rhamnopyranoside	Not found	Not found	S. macrostema	Gutierrez (2013)
5,3′,4′,5′-Tetrahydroxyflavanone-7-O-β-d-rhamnopyranoside	Not found	Not found	S. macrostema	Gutierrez (2013)
5,3′,4′,5'-Tetramethoxyflavanone-7-O-β-d-rhamnopyranoside	Not found	Not found	S. macrostema	Gutierrez (2013)
5,4′-Dimethoxy-3′-hydroxyflavone-7-β-d-rhamnopyranoside	Not found	Not found	S. macrostema	Gutierrez (2013)
Flavanone glucosides
Hesperidin	C28H34O15	610.560608	S. aintabensis	Askun et al. (2013)
			S. cilicica	Cetojevic-Simin et al. (2012)
			S. corulea	Cetojevic-Simin et al. (2012)
			S. icerica	Cetojevic-Simin et al. (2012)
			S. mexicana	Estrada-Reyes et al. (2008)
			Naringin	C27H32O14	580.534607	S. aintabensis	Askun et al. (2013)
S. cilicica	Cetojevic-Simin et al. (2012)
S. corulea	Cetojevic-Simin et al. (2012)
S. icerica	Cetojevic-Simin et al. (2012)
Naringenin glucoside	C21H22O10	434.393402				S. hortensis	Kosar et al. (2005)
Prunin	C21H22O10	434.393402	S. macrostema	Gutierrez (2013)
Flavone glucosides
Vitexin	C21H20O10	432.377502	S. cilicica	Cetojevic-Simin et al. (2012)
			S. corulea	Cetojevic-Simin et al. (2012)
			S. icerica	Cetojevic-Simin et al. (2012)
			Apigenin-7-glucoside	C21H20O10	432.377502	S. cilicica	Cetojevic-Simin et al. (2012)
S. corulea	Cetojevic-Simin et al. (2012)
S. icerica	Cetojevic-Simin et al. (2012)
Luteolin-glucoside	C21H20O11	448.376892				S. hortensis	Kosar et al. (2005)
Apigenin glucoside	C21H20O10	432.377502	S. hortensis	Kosar et al. (2005)
Flavonol glucosides
Rutin	C27H30O16	610.517517	S. cilicica	Cetojevic-Simin et al. (2012)
			S. corulea	Cetojevic-Simin et al. (2012)
			S. icerica	Cetojevic-Simin et al. (2012)

In this section, flavonoids are presented in the table by dividing into several sub-groups such as flavones, flavonols, flavanones, flavan-3-ols, and flavanonols. Glucosides are also presented such as flavanoid glucosides, flavanone glucosides, flavone glucosides, and flavonol glucosides.

As can be seen from the table, luteolin and apigenin are the most common flavones found in many Satureja species. Studies reporting these phytochemicals are particularly focused on S. hortensis. These are followed by 5,6-dihydroxy-7,8,3′,4′-tetramethoxyflavone, 5,6,3′-trihydroxy-7,8,4′-trimethoxyflavone, and thymonin (5,6,4′-trihydroxy-7,8,3′-trimethoxyflavone). All three compounds are found in S. atropatana (Gohari et al., 2009; Moradi et al., 2008).

According to the literature survey, five different flavonols (quercetin, epicatechin-3-O-gallate, epigallocatechin-3-O-gallate, kaempferol, and myrcetin) were determined in Satureja species. As presented in Table 5, the most common is quercetin which is determined in S. aintabensis P. H. Davis, S. cilicica, S. corulea L., S. hortensis, S. icerica, S. montana, and S. sahendica (Alizadeh, 2011; Cetojevic-Simin et al., 2012; Exarchou et al., 2002; Kayedi et al., 2014).

Flavan-3-ols (catechin and epicatechin) (Cetkovic et al., 2007b; Cetojevic-Simin et al., 2012; Palavra et al., 2011; Silva et al., 2009) and flavanonols (aromadendrin, taxifolin) (Malmir et al., 2012; Skoula et al., 2005) are other flavonoid aglycones found in Satureja species (Table 5).

Apart from the flavonoid aglycones, glucoside derivatives of flavonoids have also been investigated several times. According to the literature survey; flavonoid glucosides, flavanone glucosides, flavone glucosides, and flavonol glucosides are the main chemical groups identified in Satureja species (Table 5).

Gutierrez (2013) has determined six new flavonoid glucosides [5-hydroxy-3,6,4′-trimethoxyflavonol-7-C-α-l-rhamnopyranosyl-(1 → 3)-β-d-glucopyranoside (1), 4′-methoxy-5,7,3′,5′-tetra-hydroxyflavanone-3-O-β-d-rhamnopyranosyl-(1 → 2)-β-d-rhamnopyranoside (2), 5,4′-dimethoxy-7,3′,5′-trihydroxyflavanone-3-O-β-d-rhamnopyranoside (3), 5,3′,4′,5′-tetrahydroxyflavanone-7-O-β-d-rhamnopyranoside (4), 5,3′,4′,5′-tetramethoxyflavanone-7-O-β-d-rhamnopyranoside (5), and 5,4′-dimethoxy-3′-hydroxyflavone-7-β-d-rhamnopyranoside (6)], and a flavanone glucoside (prunin) in S. macrostema Bentn.

On the other hand, Cetojevic-Simin et al. (2012) have also identified flavanone glucosides (naringin, naringenin glucoside, and hesperidin), flavone glucosides (vitexin, apigenin-7-glucoside, luteolin glucoside, and apigenin glucoside), and rutin (a flavonol glusocide) in S. cilicica, S. corulea, and S. icerica. Compounds identified in Satureja species are presented in Table 5 in detail with their reference data.

Other phytochemicals

As far as the literature survey could ascertain, apart from the terpenoids, phenolic acids, and flavonoids, some other phytochemicals have also been reported several times. Data obtained from this survey are presented in Table 6.

Table 6. Other compounds of Satureja species.

Compound	Plant Species	Reference
Fatty acids
Linoleic acid	S. adamovicii	Kitic et al. (1999)
	S. biflora	Matasyoh et al. (2007)
	S. fukarekii	Kitic et al. (1999)
	Linolenic acid	S. adamovicii	Kitic et al. (1999)
S. fukarekii		Kitic et al. (1999)
Palmitic acid		S. adamovicii	Kitic et al. (1999)
	S. fukarekii	Kitic et al. (1999)
	Oleic acid	S. adamovicii	Kitic et al. (1999)
S. fukarekii		Kitic et al. (1999)
9-Octadecenoic acid methyl ester		S. cuneifolia	Goren et al. (2003)
	S. thymbra	Goren et al. (2003)
	Hexadecanoic acid methyl ester (methyl palmitate)	S. cuneifolia	Goren et al. (2003)
S. thymbra		Goren et al. (2003)
9,12,15-Octadecatrienoic acid methyl ester (Z, Z, Z)		S. cuneifolia	Goren et al. (2003)
	S. thymbra	Goren et al. (2003)
	Octadecanoic acid methyl ester	S. cuneifolia	Goren et al. (2003)
S. thymbra		Goren et al. (2003)
Enzymes
Flavocytochrome b(2)	S. hortensis	Arzi et al. (2010)
Cytochrome b(2)  (NAD-independent lactate dehydrogenase)	S. hortensis	Arzi et al. (2010) and Arzi et al. (2008)
Peroxidase isoenzymes	S. hortensis	Keyhani et al. (2009)
o-Dianiside peroxidases	S. hortensis	Paknia et al. (2006)
Lignin peroxidase	S. hortensis	Keyhani and Keyhani (2006)
Ascorbate peroxidase	S. hortensis	Keyhani and Keyhani (2006)
Superoxide dismutase	S. hortensis	Keyhani and Keyhani (2006)
Tocopherols
α-Tocopherol	S. thymbra	Lagouri and Boskou (1996)
β-Tocopherol	S. thymbra	Lagouri and Boskou (1996)
γ-Tocopherol	S. thymbra	Lagouri and Boskou (1996)
δ-Tocopherol	S. thymbra	Lagouri and Boskou (1996)
Triterpene acids
Ursolic acid	S. atropatana	Moradi et al. (2008)
	S. khuzestanica	Moghaddam et al. (2007)
	S. mexicana	Estrada-Reyes et al. (2008)
	S. montana	Razborsek et al. (2008a)
	S. parvifolia	van Baren et al. (2006)
	S. spicigera	Gohari et al. (2012)
	Oleanolic acid	S. atropatana	Moradi et al. (2008)
S. montana		Razborsek et al. (2008a)
S. parvifolia		van Baren et al. (2006
S. spicigera		Gohari et al. (2012
Betulinic acid		S. montana	Razborsek et al. (2008a, 2008b)
Phytosterols
β-Sitosterol	S. atropatana	Moradi et al. (2008)
	S. khuzestanica	Moghaddam et al. (2007)
	S. mexicana	Estrada-Reyes et al. (2008)
	S. spicigera	Gohari et al. (2012)
	β-Sitosterol-3-O-beta-d-glucopyranoside	S. khuzestanica	Moghaddam et al. (2007)
Alkana Homologues
Nonacosane	S. adamovicii	Kitic et al. (1999)
	S. fukarekii	Kitic et al. (1999)
	Hentricontane	S. adamovicii	Kitic et al. (1999)
S. fukarekii		Kitic et al. (1999)
Dihydrochalcone
Nubigenol	S. atropatana	Moradi et al. (2008)
	S. spicigera	Gohari et al. (2012)
	Erigeroside	S. khuzestanica	Tafazzoli et al. (2008)

As can be seen from the table, compounds under this topic are divided into several groups as the following: fatty acids, enzymes, tocopherols, triterpene acids, phytosterols, and alkane homologues in addition to nubigenol (dihydrochalcone) and erigeroside.

The vast majority of studies reported under this topic have especially focused on fatty acids. Kitic et al. (1999) reported linoleic, linolenic, palmitic, and oleic acids in S. adamovicii Silic and S. fukarekii Silic. Additionally, Goren et al. (2003) have reported fatty acids of S. cuneifolia and S. thymbra (9-octadecenoic acid methyl ester, hexadecanoic acid methyl ester (methyl palmitate), 9,12,15-octadecatrienoic acid methyl ester (Z, Z, Z), and octadecanoic acid methyl ester).

Accoring to the literature survey, only S. hortensis was evaluated for its enzyme content so far. Flavocytochrome b(2), cytochrome b(2) (NAD-independent lactate dehydrogenase), peroxidase isoenzymes, o-dianiside peroxidases, lignin peroxidase, ascorbate peroxidase, and superoxide dismutase are the enzymes identified in this species (Arzi et al., 2008, 2010; Keyhani & Keyhani, 2006; Keyhani et al., 2009; Paknia et al., 2006).

Data concerning tocopherols (α-, β-, γ-, δ-tocopherols), triterpene acids (ursolic, oleanolic, and betulinic acids), phytosterols (β-sitosterol and β-sitosterol-3-O-β-d-glucopyranoside), alkana homologues (non-acosane and hentricontane), nubigenol (a dihydrochalcone), and erigeroside are also presented in Table 6 with their references.

Inorganic compounds

In addition to organic phytochemicals given above, inorganic compounds of Satureja species have also been reported elsewhere. Dunkic et al. (2012) have reported Ca, Mg, K, Na, and Hg, contents of S. montana and S. subspicata (Bartl. ex Vis). According to this report, high quantities of Ca and Mg, moderate concentrations of K and Na and low concentrations of Hg were determined in these species.

Mumivand et al. (2010) have reported the effect of nitrogen and CaCO₃ on mineral content. According to this report, nitrogen fertilization increased N, P, and B contents. K and Mg were also affected by nitrogen fertilization. Additionally, Ca, Fe, Mn, Zn, and Cu were not affected by nitrogen. However, CaCO₃ increased Ca and B contents while decreasing P, Mg, Fe, Mn, and Zn contents. According to the authors, Zn was not affected by CaCO₃.

Grzeszczuk and Jadczak (2009) have reported total nitrogen, total protein, total sugar, reducing sugar, saccharose, chlorophyll a, chlorophyll b, vitamic C, nitrat, and nitrit concentrations of S. hortensis. Satureja abyssinica (Benth.) Briq. have also been reported to have a high concentration of chromium by Odhiambo and Howarth (2002).

Conclusion

In this paper, as can be seen from the section given above, Satureja species were evaluated in terms of its pharmacological and phytochemical characteristics. In the paper, all attempts were made to place all articles published on this subject, as much as possible. However, there may be some articles possibly overlooked. In this review, phytochemical compositions of species were evaluated in details. On the contrary, due to the page limitations and intensity of data in relevant area, some parts were necessarily written down as summary. As can be seen from the “Pharmacology” section, antimicrobial and antioxidant activity potentials of Satureja species could not be presented in full detail. Additionally, under “Other activities” title, apart from the activities given in details, many reports could be noted only in the form of title. We hope that the missing points of this review encourage other scientists to gather all the required information in this field.

Declaration of interest

The authors report no conflicts of interest. The authors alone are responsible for the content and writing of this article.