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Plants of the Lamiaceae family are important ornamental, medicinal, and aromatic plants, many of which produce essential oils that are used in traditional and modern medicine, and in the food, cosmetics, and pharmaceutical industry. Various species of the genera Hyssopus, Leonurus, Mentha, Nepeta, Origanum, Perovskia, Phlomis, Salvia, Scutellaria, and Ziziphora are widespread throughout the world, are the most popular plants in Uzbek traditional remedies, and are often used for the treatment of wounds, gastritis, infections, dermatitis, bronchitis, and inflammation. Extensive studies of the chemical components of these plants have led to the identification of many compounds, as well as essentials oils, with medicinal and other commercial values. The purpose of this review is to provide a critical overview of the literature surrounding the traditional uses, ethnopharmacology, biological activities, and essential oils composition of aromatic plants of the family Lamiaceae, from the Uzbek flora.
The Republic of Uzbekistan is located in the center of Eurasia. About 85% of its territories are deserts, and about 15% are mountains and foothills. The Uzbek flora accounts for approximately 4350 species of vascular plants, including large numbers of endemic, endangered, and globally important species. Plants endemic to Uzbekistan constitute 20% of all plants; and a majority of these grow in the mountains. The floristic data for several regions of Uzbekistan is imperfect, and studies are continuing [1,2].
One of the famous medicinal aromatic plant families is the Lamiaceae family (alt. Labiatae), also known as the mint family. Aromatic medicinal plants from this family have long been used in Uzbek traditional medicine. The aim of this review is to present a critical overview of the ethnopharmacology, ethnobotany, phytochemistry, essential oils composition, and biological activities of medicinal plants of the family Lamiaceae, e.g., Hyssopus seravschanicus, Leonurus panzerioides, L. turkestanicus, Mentha longifolia var. asiatica, Nepeta alatavica, N. olgae, Origanum tyttanthum, Perovskia scrophulariifolia, Phlomis thapsoides, Salvia korolkovii, S. sclarea, Scutellaria immaculata, S. ramosissima, S. schachristanica, Ziziphora clinopodioides, and Z. pedicellata, from the Uzbek flora. This review has been compiled using references from major databases, such as Science Direct, SciFinder, Pubmed, and Google Scholar Databases. The search included articles published to date.
The genus Hyssopus L. consists of over 15 species worldwide. Hyssopus is a source of volatile oils and its constituents are mostly sesquiterpenes, bicyclic monoterpenes, and some acids . Only one species of this genus, Hyssopus seravschanicus, grows in Uzbekistan . Hyssopus seravschanicus Pazij is a perennial, branched, semi-shrub that is native to the the Republic of Uzbekistan. Several researchers have studied the essential oils content of H. seravschanicus [3,4,5]. The most abundant compounds identified in the essential oils of H. seravschanicus, were pinocamphone (71.0%), β-pinene (8.6%), 1,8-cineole (6.4%), carvacrol (1.6%), cis-ocimene (1.4%), p-cymene (1.3%) and sabinene (1.3%) (Figure 1) (Table 1) .
The genus Leonurus L. (subfamily Lamioideae) comprises 25 species . The Leonurus are characterized by the presence of iridoid glycosides and a lower content of essential oils. This genus is represented in Uzbekistan by four species. Leonurus panzerioides Popov is a perennial shrub that grows in Western Tien Shan and the Pamir-Alay mountains, on stony and gravelly slopes [2,7]. A tincture of the herb of L. panzerioides has been known to possess a sedative effect, which is twice as strong as the effect of a valerian tincture (Table 1) . The main constituents of the essential oils of L. panzerioides were found to be eugenol (30.93%), p-vinyl guaiacol (15.77%), dihydroactinidiolide (8.95%), phenyl ethyl alcohol (6.51%), verbenone (5.83%), and p-cymen-8-ol (5.24%). Twenty-four compounds were identified in the oil of L. panzerioides, which accounted for 99.98% of the total oil .
Leonurus turkestanicus V. I. Krecz & Kuprian is a perennial shrub that grows in the plains and highlands of Asia on stony, shallow-soiled slopes, floodplains, streamsides, and among trees and other shrubs . A decoction of the above-ground parts is used to treat various ailments of the heart, stomach, and nervous system [8,11]. Previous phytochemical investigations of the aerial parts of L. turkestanicus identified flavonoids, iridoids, alkaloids, and fatty acids. Thirty-nine chemical constituents were detected by GC–MS analysis of the essential oils of L. turkestanicus, representing 99.98% of total oil components. The essential oil from the aerial parts of L. turkestanicus had oxygenated monoterpenoids and sesquiterpenoids as the major components, and thus shared the characteristics of thymol chemotype. The principal constituents of the essential oils of this species were found to be thymol (40.10%), octen-3-ol (13.07%), carvacrol (5.83%), and β-caryophyllene (5.61%) (Table 1) .
Mentha L. (mint) is a well-known genus due to its medicinal and aromatic value. It is represented by about 19 species and 13 natural hybrids, mainly perennial herbs, which grow wildly in damp or wet places throughout the temperate regions of Europe, Asia, Africa, Australia, and North America. Species of the genus Mentha have been reported to contain a range of components, including cinnamic acids, flavonoids, and steroidal glycosides. However, the main active component of the genus Mentha is essential oil, which is reported to govern its various properties . Three species of the Mentha grow in Uzbekistan . Mentha longifolia var. asiatica (Boriss) Rech. f., is commonly used as a cooking herb by the people. It has a pleasant taste and is a popular flavouring for food and drink. Thirty-seven compounds were characterized from M. longifolia var. asiatica, representing 97% of the total components detected. The major constituents of the oil were found to be trans-piperitone oxide (64.51%), piperitenone oxide (12.34%), cis-piperitone oxide (7.24%), thymol (2.60%), and spathulenol (2.36%) (Table 1) .
The genus Nepeta L. comprises perennial or annual herbaceous, small shrubs, and rarely includes trees. It is comprised of more than 200 species. This genus has a widespread distribution in the temperate regions of Asia, Europe, and North Africa. Some Nepeta species are widely used in traditional medicine, due to their diuretic, antispasmodic, anti-asthmatic, febrifuge, emmenagogue, sedative, and antiseptic properties . In Uzbekistan, the genus Nepeta is represented by 19 species . Nepeta alatavica Lipsky is a perennial plant that grows in the Tien Shan mountains (Kyrgyz Alatau ridges, Talas Alatau, Karzhantau, Ugam, Pskem, Chatkal mountains). The major constituents of the essential oils of N. alatavica were thymol (48.5%), carvacrol (7.5%), verbenone (7.7%), and 1-octen-3-ol (4.1%) (Table 1) .
In Uzbekistan, Nepeta olgae Regel grows as an aromatic perennial plant in the foothills and lowlands of the Syrdaya region, Kyzylkum, and the Surkhan-Sherabad and Ferghana Valleys . The leaves of this plant have the richest odour intensity, and are used primarily for its fragrance. The N. olgae oil is dominated by acetylcyclohexene (31.5%), 4-tridecyne (13.2%), 2-methyl cyclopentanone (6.8%), and 1,8-cineole (6.0%) (Table 1) .
The genus Origanum L. consists of 43 species and 18 hybrids; most of which are distributed through the eastern Mediterranean region . Only one species, Origanum tyttanthum Gontsch., is found in Uzbekistan. This herbaceous perennial, rhizomatous plant, grows on rocky and pebbly slopes. The plant contains phenolic glycosides, lipids, and coumarins . Forty compounds were characterized, representing 98.6% of the total components in its essential oils . The major components of the essential oils were reported to be carvacrol (42.76%), thymol (27.18%), γ-terpinene (9.50%), p-cymene (5.90%), and β-bisabolene (2.65%) . In a different study , O. tytthanthum oils were analyzed to reveal about 30 compounds, 20 of which were identified, and the major ones were thymol and carvacrol (48%–89%). The above-ground part of O. tyttanthum was found to contain 0.3%–2.1% oil. The oil content in the different parts of this plant changes considerably, depending on the different conditions: phase of vegetation, sunlight exposition, and altitude of plant growth (Table 1).
The genus Perovskia Kar. is made up of seven different species , four of which grow in Uzbekistan . Perovskia scrophulariifolia Bunge is an aromatic sub-shrub featuring extremely branched stems, paniculate leaves, and small flowers. It is mainly known for its ornamental and flavouring qualities. From the aerial parts of P. scrophulariifolia, abietane-type diterpenoids, flavone glycosides, anthocyanins, and coumarins were isolated and identified . Constituents of the essential oil of the aerial parts of P. scrophulariifolia growing in Turkmenistan, were reported as being borneol, camphene, geraniol, linalool, α-pinene, sabinene, terpinene, and terpinolene, without quantitative indication . In another study, the essential oil of P. scrophulariifolia  revealed the presence of a total of at least 20 components. Farnesene and the products of its oxidation, and some modifications of sesquiterpene hydrocarbons with a naphthalene nucleus, were identified. Of the monoterpenoids, bornyl acetate predominated, and of the sesquiterpenoids, α- and β-caryophyllenes were identified. In a separate study, major constituents found in the oils P. scrophulariifolia were as follows: 1,8-cineole (11.0%), caryophyllene oxide (10.0%), camphor (9.0%), humulene epoxide II (7.9%), bornyl acetate (7.8%), and p-cymene (5.7%) (Table 1) .
The genus Phlomis L. consists of 75 species of perennial shrubs, occurring from the Mediterranean to Central Asia. In Uzbekistan, the genus Phlomis is represented by 15 species. Phlomis thapsoides Bunge is a perennial herb growing wild in western Pamir-Alay. The aerial parts of this species are used in some areas of Asia for feeding animals, and to dye wool and silk . Previous phytochemical studies of the aerial parts of P. thapsoides afforded iridoids, sterols, aliphatic ketones, and essential oils . The GLC-MS analysis of the essential oil obtained from the aerial parts of P. thapsoides, identified phenylethyl alcohol (6.81%), trans-3-hexenol (5.55%), 1-octen-3-ol (5.10%), α-cadinol (4.92%), α-muurolol (4.67%), and linalool (3.69%) as the main volatile constituents (Table 1).
The Salvia L. (sage) belongs to the Mentheae Tribe, and is the largest and most diverse genus of the Lamiaceae. This genus contains over 900 species, throughout the world. Many species of the Salvia have been used worldwide as a flavouring agent, as well as in traditional herbal medicine. A total of 21 species of the Salvia, including Salvia korolkovii Regel et Schmalh., are native to Uzbekistan [2,25]. This plant is endemic to Uzbekistan, and one of the most important aromatic plants and sources of essential oils. The main constituents of the essential oils of S. korolkovii were found to be 1,8-cineole (29.3%), camphor (9.8%), β-caryophyllene (8.5%), bornyl acetate (7.7%), caryophyllene oxide (7.2%), borneol (5.6%), camphene (3.4%), and limonene (3.3%) (Table 1) .
Salvia sclarea L. (clary sage) is a well known aromatic plant, from which an oil used to be produced in large quantities in the former Soviet Union. Apart from the various medicinal uses, the essential oils of clary sage are widely applied in the food and cosmetic industries, wine making. and as a tobacco flavouring agent . In addition, some authors indicated that the oil composition of S. sclarea was affected by the method of isolation, as well as by the plant origins and organs used for oil isolation [28,29,30]. The principal components of the oils of S. sclarea collected from Uzbekistan were reported to be linalool (22%–32%), linalyl acetate (25%–51%) α-terpineol (10.0%–11.0%), and geranylacetate (5.4%–6.7%) (Table 1) .
The genus Scutellaria L. includes about 350 species, commonly known as skullcaps , 38 species of which grow in Uzbekistan . Scutellaria immaculata Nevski ex Juz. is a semi-shrub featuring beautiful white flowers, and grows on rocky and gravelly slopes. The flavonoids of this species have been well studied . Constituents of the essential oil of the aerial parts of S. immaculata growing in Uzbekistan were reported as including acetophenone (30.39%), eugenol (20.61%), thymol (10.04%), linalool (6.92%), 1-octen-3-ol (2.89%), 4-vinylguaicol (2.50%), and 1,8-cineol (2.25%) (Table 1) .
Scutellaria ramosissima Popov is native to Uzbekistan and grows in Northern Tien Shan, Pamir-Altay mountains (Central Asia), on the midlands of the high-altitude belt. In Uzbek folk medicine, water extracts (tea and infusion) of S. ramosissima are widely applied for epilepsy, inflammation, allergies, and nervous tension. The bioactivity of the plant is most likely due to the major components: flavonoids. The chloroform extract of S. ramosissima showed potent cytotoxic effects to Trypanosoma brucei TC221, and HeLa, HepG-2, and MCF-7 cancer cells . The essential oils from S. ramosissima showed that germacrene D (23.96%), β-caryophyllene (11.09%), linalool (9.63%), hexadecanoic acid (8.34%), caryophyllene oxide (5.90%), eugenol (5.29%), acetophenone (4.67%), thymol (3.01%), and 4-vinylguaicol (2.42%), were the principal components (Table 1) .
Scutellaria schachristanica Juz is a perannial that grows in the rocky and gravelly slopes of the highlands of Pamir-Alai (ridges Turkestan and Zeravshan). Several flavonoids were isolated from the aerial parts of this plant. The methanol/water extracts of Scutellaria are known as potent-free radical scavengers . However, the essential oils obtained from the aerial parts of S. schachristanica exhibited weaker antioxidant effects in the DPPH, ABTS, and FRAP assays. The oil of the Uzbek S. schachristanica was reported to be largely composed of acetophenone (34.74%), linalool (26.98%), eugenol (20.67%), 1-octen-3-ol (3.73%), β-terpineol (3.57%), 2-methoxy-p-cresol (1.89%), and 4-vinylguaicol (1.64%) (Table 1) .
The species of the genus Ziziphora are annuals or perennials, and herbaceous or sub-shrubs. The world population of this genus is represented by more than 30 different species. The Ziziphora species are rich in flavonoids, caffeic acid derivatives, fatty acids, triterpenes, and sterols. The essential oils of Ziziphora have been well studied . In the flora of Uzbekistan, this genus consists of eight species, including Ziziphora capitata L., Z. clinopodioides Lam., Z. interrupta Juz., Z. pamiroalaica Juz., Z. pedicellata Pazij & Vved., Z. persica Bunge, Z. suffruticosa Pazij & Vved., and Z. tenuior L. .
Ziziphora clinopodioides Lam. is an important aromatic, edible medicinal plant. The leaves, flowers, and stem of the plant are frequently used as a wild vegetable, or as an additive to foods. The chemical constitutions of Z. clinopodioides growing in Iran, Turkey, Kazakhstan, Altai Republic (Russia), Tajikistan, and Urumqi (China), have been previously analyzed . The compositions of the samples of the essential oils from the inflorescences and leaves collected in Uzbekistan, were determined . The main component of the essential oils of the inflorescences and leaves was identified as pulegone, a substance that is characteristic of the species which were previously investigated. The amount of pulegone in the inflorescences was 88%, and was 75% in the leaves ((in leaves: pulegone (75.0%), menthone (9.6%), menthol (4.7%), caryophyllene (1.5%), pinocamphone (1.5%), limonene (1.4%); in inflorescences: pulegone (88.0%), menthone (3.2%), limonene (1.6%), pinocamphone (1.1%), caryophyllene (0.9%), menthol (0.5%)). The essential oil of the inflorescences differed from that of the leaves, by the presence of α-pinene, sabinene, and myrcene, and also by the absence of substances such as linalool, linalyl acetate, carvacrol, and α-cedrene, that were present in the essential oil of the leaves.
Ziziphora pedicellata Pazij Vved. grows on the stony, gravelly slopes of the foothills and midlands of Tien Shan. This plant contains organic acids, essential oil, vitamin C, saponins, and flavonoids. The seeds and leaves contain carotenoids and lipids. The flowers contain terpenes [8,37]. The essential oils obtained by hydro-distillation were analyzed, and a total of 31 compounds were identified from Z. pedicellata. The essential oils of Z. pedicellata were rich in the oxygenated monoterpenes pulegone (62.0%), isomenthone (11.5%), menthol (9.2%), menthone (5.5%), and β-pinene (1.0%) (Table 1) .
Plant species from the Lamiaceae family have been used in herbal medicine for thousands of years. Traditional applications of the Lamiaceae family show high applicability as a common tea, flavours, insect repellant, in flu control, and as an anti-inflammatory, sedative, and analgesic. Mainly essential oils, terpenoids, phenolic compounds, flavonoides, and iridoids, have been reported from the members of this family. Many of the medicinal uses are presumed to be connected to the terpenic constituents of the essential oils of these plants.
Table 1 presents the various ethnopharmacological uses of the Lamiacea species, that have been widely used in Uzbek traditional medicine. The local names, as well as the collection location, growing environment, and their essential oil composition, are also included.
The present review provides, for the first time, an updated compilation of the documented ethnopharmacological information in relation to the ethnomedicinal, ethnobotanical, phytochemistry, and biological activities of 16 aromatic and medicinal plants from the Lamiaceae family of the Uzbek flora. Information on their traditional medicinal uses, and the compounds identified in the essential oils obtained from Hyssopus seravschanicus, Leonurus panzerioides, L. turkestanicus, Mentha longifolia var. asiatica, Nepeta alatavica, N. olgae, Origanum tyttanthum, Perovskia scrophulariifolia, Phlomis thapsoides, Salvia korolkovii, S. sclarea, Scutellaria immaculata, S. ramosissima, S. schachristanica, Ziziphora clinopodioides, and Z. pedicellata, has also been presented. The available literature showed that most of the bioactivities and medicinal properties of these species could be attributed to their essential oils, which contain a variety of functional bioactive compounds, known to have applications in the food, feed, pharmaceutical, and cosmetic industries.
Nilufar Z. Mamadalieva thanks UNESCO-L’ORÉAL and OWSD-TWAS-Elsevier for generous financial supports.
N.Z.M. and S.S.A. conceived and organized the review; D.K.A., A.T., E.O., L.N., and S.D.S. contributed to the writing and editing of the manuscript.
The authors declare no conflict of interest.