Caryologia. International Journal of Cytology, Cytosystematics and Cytogenetics 74(4): 21-28, 2021 Firenze University Press www.fupress.com/caryologia ISSN 0008-7114 (print) | ISSN 2165-5391 (online) | DOI: 10.36253/caryologia-641 Caryologia International Journal of Cytology, Cytosystematics and Cytogenetics Citation: Halil Erhan Eroğlu, Esra Mar- tin, Ahmet Kahraman, Elif Gezer Aslan (2021) The new chromosomal data and karyotypic variations in genus Salvia L. (Lamiaceae): dysploidy, polyploidy and symmetrical karyotypes. Caryolo- gia 74(4): 21-28. doi: 10.36253/caryolo- gia-641 Received: October 01, 2019 Accepted: September 24, 2021 Published: March 08, 2022 Copyright: © 2021 Halil Erhan Eroğlu, Esra Martin, Ahmet Kahraman, Elif Gezer Aslan. This is an open access, peer-reviewed article published by Firenze University Press (http://www. fupress.com/caryologia) and distributed under the terms of the Creative Com- mons Attribution License, which per- mits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Data Availability Statement: All rel- evant data are within the paper and its Supporting Information files. Competing Interests: The Author(s) declare(s) no conflict of interest. ORCID HEE: 0000-0002-4509-4712 EM: 0000-0002-5484-0676 AK: 0000-0002-9344-1993 EGA: 0000-0003-4645-3892 The new chromosomal data and karyotypic variations in genus Salvia L. (Lamiaceae): dysploidy, polyploidy and symmetrical karyotypes Halil Erhan Eroğlu1,*, Esra Martin2, Ahmet Kahraman3, Elif Gezer Aslan4 1 Department of Biology, Faculty of Science and Art, Yozgat Bozok University, Yozgat, Turkey 2 Department of Biotechnology, Faculty of Science, Necmettin Erbakan University, Konya, Turkey 3 Department of Biology, Faculty of Science and Arts, Uşak University, Uşak, Turkey 4 Department of Medical Services and Techniques, Vocational School of Health Services, Kırklareli University, Kırklareli, Turkey *Corresponding author. E-mail: herhan.eroglu@bozok.edu.tr Abstract. In this study, it was aimed to determine the chromosome number of 21 Sal- via L. species, to determine chromosome morphology, to reveal karyotype analysis in detail and to contribute to the cytotaxonomy of Salvia. In this context, the results are as follows: (i) the first report for the number of chromosomes of ten species, namely S. corrugata Vahl. (2n = 16), S. curviflora Benth. (2n = 16), S. darcyi J.Compton, S. greg- gii A.Gray, S. longifolia Nutt., S. vitifolia Benth. (2n = 22), S. subrotunda A.St.-Hil. ex Benth. (2n = 44), S. oppositiflora Ruiz & Pav. (2n = 56), S. stolonifera Benth. and S. atrocyanea Epling (2n = 60); (ii) the karyotypic variations and new chromosome num- bers different from previous reports for three species, namely S. cardiophylla Benth. (2n = 36), S. cuspidata Ruiz & Pav. (2n = 44) and S. microphylla Sessé & Moc. (2n = 46); (iii) the same chromosome numbers from previous reports for eight species, namely S. campanulata Wall. ex Benth. (2n = 16), S. elegans Vahl. (2n = 20), S. involucrata Cav., S. mexicana Sessé & Moc. (2n = 22), S. apiana Jeps., S. leucophylla Greene, S. mellifera Greene (2n = 30), and S. splendens Ker Gawl. (2n = 44); (iv) the detailed chromosome measurements and karyotype analyses for all species studied for the first time; (v) the symmetrical karyotypes for all studied species; (vi) the variations resulting from dys- ploidy or polyploidy and discussing their reasons. Keywords: chromosomal alteration, karyotype asymmetry, sage, Turkey. INTRODUCTION The word Salvia that means sage in Turkish is derived from the Latin salvare, which means protect and heal because of its medicinal properties. The genus Salvia is placed in the family Lamiaceae and is one of the largest 22 Halil Erhan Eroğlu et al. genera of the family with nearly 1000 perennials, bienni- al or annual, often strongly aromatic species throughout the world (Sheidai and Alijanpoo 2011). This ratio cor- responds to one quarter of the family. The Salvia species usually spread in tropical and temperate regions of the world. The species are mostly distributed in three differ- ent regions: Central and South America (about 500 spe- cies), West Asia (about 200 species) and East Asia (about 100 species) (Walker and Sytsma 2007). Turkey, which has 98 species in terms of the diversity of Salvia species, is an important gene center in Asia (Hedge 1982; Kahra- man et al. 2011). Aboveground organs of Salvia species have been used in cough, colds, teeth, stomach and abdominal pains and skin diseases since ancient times. Most of Sal- via species are used as folk medicine because of their antioxidant, antidiabetic, antimicrobial, antitumor, anti- plasmodial, antihypertensive and anti-inf lammatory properties (Ulubelen 2003; Kamatou et al. 2008; Şenol et al. 2010). Some Salvia species have been reported to be used to prevent memory loss (Perry et al. 1996). In addi- tion, Salvia species are frequently used in food, perfum- ery, cosmetics and pharmaceutical industries (Chalchat et al. 1998; Baylac and Racine 2003). Many Salvia spe- cies are easily cultured frequently due to their aromatic nature; and because of their beautiful appearance, they are grown as decorative ornamental plants in parks and gardens (Nakipoğlu 1993; Marin et al. 1996). Many karyological reports showed that Salvia is a polybasic genus with diverse chromosome numbers in different regions of the world and the species are polyploid origins (Sheidai and Alijanpoo 2011). It was reported that the basic number is x = 16 for California species (Epling et al. 1962); is x = 11 for species of Rus- sia and Europe (Patudin et al. 1975); is x = 7 for Medi- terranean species (Afzal-Rafii 1976). According to the chromosome databases, comprehensive chromosomal reports exist in genus Salvia. Due to the high number of species and samples, there may be some cytotaxonomic uncertainties. The purpose of this work is to contribute to the cytotaxonomy of Salvia with the following ques- tions: (1) The chromosome numbers of which species will be reported for the first time? (2) Are there spe- cies with karyotypic variations and new chromosome numbers different from previous reports? (3) What are the detailed chromosome measurements and karyotype analysis results for all species? (4) What are the karyo- type asymmetry states for all species? Symmetrical or asymmetrical. (5) What are the chromosomal variations caused by polyploidy and dysploidy in genus Salvia? (6) What are the possible causes of polyploidy, dysploidy, and symmetrical/asymmetrical karyotypes? MATERIALS AND METHODS The seeds of the plants included in the study were provided by Mr. Robin Middleton, who cultivated many Salvia species in his personal botanical garden in Eng- land. Identification and confirmation of the specimens were performed by the third author of this study. The cytogenetical study was conducted on root tips germinated on wet filter paper in Petri dishes. After ger- mination, the fresh root tip meristems were pretreated in α-mono-bromonaphthalene at 4°C for 16 hours, fixed in glacial acetic acid and absolute alcohol (1:3) at 4°C for 24 hours, deposited in 70% ethanol at 4°C, and then hydro- lyzed in 1 N HCl at room temperature for 12 minutes. Finally, they were squashed and stained in 2% aceto-orce- in. Permanent slides were prepared using Standard liquid nitrogen method (Altay et al. 2017; Martin et al. 2019). Karyotypes were determined using Image Analysis System (Bs200Pro) on a personal computer. 10 mitot- ic plates were assessed to determine the chromosome numbers. The following variables were measured: long arm (la), short arm (sa), total chromosome length (la + sa), arm ratio (la / sa), centromeric index [(sa / la + sa) × 100], total haploid length (THL), mean chromosome length (MCL), and relative length (RL%). Centromere positions and karyotype formulae of 17 Salvia species were determined. From the point of view of chromo- some morphology, median (M, m), submedian (sm) and subtelocentric (st) chromosome pairs were observed (Levan et al. 1964). As centromere positions of the other taxa (S. cardiophylla, S. cuspidata, S. oppositiflora, and S. atrocyanea) could not be determined, their total chro- mosome length and haploid chromosome length were measured. Intrachromosomal asymmetry and interchro- mosomal asymmetry were determined with the parame- ters of MCA (Peruzzi and Eroğlu 2013) and CVCL (Paszko 2006), respectively. The intrachromosomal asymmetry increases by shifting of centromere position from the center to the end of the chromosome. In this case there is a transition from median/submedian chromosomes to subterminal/terminal chromosomes. The interchromo- somal asymmetry depends on relative variation in chro- mosome length, namely it determines how different the chromosome lengths of a complement. Finally, a scatter diagram was drawn between MCA and CVCL. RESULTS Chromosomal data Chromosome records of 21 taxa are herein provided (Figure 1), ten of which are reported for the first time, 23The new chromosomal data and karyotypic variations in genus Salvia Figure 1. Mitotic metaphase chromosomes of Salvia: (A) S. corrugata, (B) S. campanulata, (C) S. curviflora, (D) S. elegans, (E) S. darcyi, (F) S. greggii, (G) S. involucrata, (H) S. longifolia, (I) S. vitifolia, (J) S. mexicana, (K) S. apiana, (L) S. leucophylla, (M) S. mellifera, (N) S. cardiophylla, (O) S. cuspidata, (P) S. splendens, (R) S. subrotunda, (S) S. microphylla, (T) S. oppositiflora, (U) S. stolonifera, (V) S. atrocyanea (scale bar: 10 μm). 24 Halil Erhan Eroğlu et al. three possess new chromosome numbers, and eight have the same results including previous reports. Ten differ- ent chromosome numbers (2n = 16, 18, 20, 22, 30, 36, 44, 46, 56, and 60) are also detected (Table 1). Among the studied taxa, the smallest and the largest chromo- some shapes are 0.53 μm in S. oppositiflora and 3.28 μm in S. campanulata, respectively. The smallest and the highest values of total haploid chromosome length are 9.12 μm in S. curviflora and 36.92 μm in S. stolonifera, respectively (Table 2). In addition, the detailed chromo- some measurements of all chromosome pairs are given in supplemental online material (Supplementary Tables 1–21). Basic numbers and ploidy levels There are six basic chromosome numbers within Salvia, namely x = 7 in only one species, x = 8 in two species, x = 9 in two species, x = 10 in six species, most common x = 11 in nine species, and x = 23 (probably dysploidy) in only one species. The ploidy levels are 2x (in 11 species), 3x (in three species), 4x (in four species), 6x (in two species), and 8x (in only one species) (Table 2). The monoploid ideograms generated by the basic chromosome numbers are given in Figure 2. Karyotype formula and karyotype asymmetry 17 taxa possess median (m) and submedian (sm), whereas none subtelocentric (st) chromosomes and telo- centric (t) chromosomes. Due to the uncertainty of cen- tromere positions, the karyotype formulae of four taxa are not given, namely S. cardiophylla, S. cuspidata, S. oppositiflora, and S. atrocyanea. Four different formulae are observed, namely (1) M-m, (2) m, (3) m-sm, and (4) M-m-sm. The MCA values for intrachromosomal asym- metry vary from 14.94 in S. curviflora to 26.01 in S. cor- rugata and are characterized by taxa with symmetric karyotypes consisting entirely of median and submedian chromosomes. The CVCL values for interchromosomal asymmetry vary from 10.73 in S. longifolia to 22.13 in S. mellifera (Table 2). Table 1. The chromosome counts of the investigated species in present and previous studies. Species Previous results References Present results Explanation n 2n 2n S. corrugata 16 First report S. campanulata 8 32 Saggoo and Bir 1986; Hu et al. 2016 16 Detailed measurements S. curviflora 18 First report S. elegans 10 Cherian and Kuriachan 1990 20 Detailed measurements S. darcyi 22 First report S. greggii 22 First report S. involucrata 7 22 + 0-1B Gill 1984; Alberto et al. 2003 22 Detailed measurements S. longifolia 22 First report S. vitifolia 22 First report S. mexicana 22 Palomino et al. 1986 22 Detailed measurements S. apiana 15, 16 32 Carlson and Stuart 1936; Stewart 1939 30 Detailed measurements S. leucophylla 24, 30 Stewart 1939; Epling et al. 1962 30 Detailed measurements S. mellifera 30, 32 Epling et al. 1962; Stewart 1939 30 Detailed measurements S. cardiophylla 44 + 0-1B Alberto et al. 2003 36 New count S. cuspidata 22 Alberto et al. 2003 44 New count S. splendens 8 32, 44 44 + 0-1B Carlson and Stuart 1936; Haque and Ghoshal 1980; Gill 1984; Alberto et al. 2003 44 Detailed measurements S. subrotunda 44 First report S. microphylla 11 22 Haque and Ghoshal 1980; Alberto et al. 2003 46 New count S. oppositiflora – – 56 First report S. stolonifera – – 60 First report S. atrocyanea – – 60 First report 25The new chromosomal data and karyotypic variations in genus Salvia DISCUSSION Table 1 shows the chromosome counts of the inves- tigated species in present and previous studies. The chromosome numbers are the first report for ten spe- cies, namely S. corrugata (2n = 16), S. curviflora (2n = 16), S. darcyi, S. greggii, S. longifolia, S. vitifolia (2n = 22), S. subrotunda (2n = 44), S. oppositiflora (2n = 56), S. stolonifera and S. atrocyanea (2n = 60). The chromo- some numbers are new counts different from previous reports for three species, namely S. cardiophylla (2n = 36), S. cuspidata (2n = 44) and S. microphylla (2n = 46). In literature, the chromosome numbers are 2n = 44 for S. cardiophylla, 2n = 22 for S. cuspidata and S. micro- phylla (Haque and Ghoshal 1980; Alberto et al. 2003). The chromosome numbers of the other eight species are the same as the previous reports, namely S. campanu- lata (2n = 16), S. elegans (2n = 20), S. involucrata and S. mexicana (2n = 22), S. apiana, S. leucophylla, and S. mel- lifera (2n = 30) and S. splendens (2n = 44) (Carlson and Stuart 1936; Epling et al. 1962; Haque and Ghoshal 1980; Palomino et al. 1986; Saggoo and Bir 1986; Cherian and Kuriachan 1990; Alberto et al. 2003). It is already known that genus Salvia includes dip- loids and polyploids (Carlson and Stuart 1936; Epling et al. 1962; Haque and Ghoshal 1980; Gill 1984; Alberto et al. 2003; Hu et al. 2016). With chromosome data avail- able at present, 11 species are diploids with 2n = 16, 18, 20, 22, and 46 (probably dysploidy) (c.52% of the spe- cies with available data) and 10 species are polyploids (c.48% of the species with available data). When previ- ous and current chromosomal data are compared, four species, S. campanulata, S. cuspidata, S. splendens, and S. microphylla, show both diploid and polyploid sta- tus (c.19% of the species with available data). This sug- gests that intraspecific polyploidy may be common in genus Salvia. The polyploid nature are demonstrated by the prevalence of cells with 2n = 30, 36, 44, 56, and 60 chromosomes in 10 species. Polyploidy originates by autopolyploidy mechanism (genome duplication in a species) and allopolyploidy (genome duplication with hybridization between species) and has played a major role in the speciation and evolution of higher plants (Demirci Kayıran and Özhatay 2017). The polyploidy possibly caused by glacial, climatic changes, altitude and high latitudes may have contributed to Salvia specia- Table 2. The karyological features of the studied Salvia taxa; karyotype formula (KF), shortest chromosome length (SC), longest chromo- some length (LC), relative length (RL), total haploid chromosome length (THL), mean chromosome length (MCL), centromeric index (CI), coefficient of variation of chromosome length (CVCL), mean centromeric asymmetry (MCA), median point (M), median (m), submedian (sm). Taxa KF SC (μm) LC (μm) RL (%) SC–LC THL (μm) MCL (μm) CI (min–max) CVCL MCA S. corrugata 8m + 8sm 1.55 2.44 10.58–16.66 14.65 1.83 31.55–41.38 15.84 26.01 S. campanulata 10m + 6sm 1.92 3.28 9.41–16.08 20.40 2.55 36.16–45.97 17.29 22.14 S. curviflora 2M + 16m 0.72 1.35 7.89–14.80 9.12 1.01 38.89–50.00 20.05 14.94 S. elegans 20m 1.21 2.16 7.48–13.35 16.18 1.62 37.13–45.22 17.06 18.55 S. darcyi 18m + 4sm 1.17 1.84 7.49–11.78 15.62 1.42 27.17–48.51 12.91 16.30 S. greggii 2M + 14m + 6sm 0.84 1.60 6.87–13.08 12.23 1.11 32.71–50.00 20.28 19.59 S. involucrata 2M + 14m + 6sm 1.03 1.76 7.05–12.04 14.62 1.33 30.00–50.00 16.00 20.90 S. longifolia 2M + 14m + 6sm 1.09 1.60 7.14–10.48 15.26 1.39 26.25–50.00 10.73 22.59 S. vitifolia 14m + 8sm 1.22 2.21 6.66–12.06 18.33 1.67 31.15–43.95 17.33 21.48 S. mexicana 20m + 2sm 0.97 1.70 6.37–11.16 15.23 1.38 35.37–44.88 15.94 17.32 S. apiana 28m + 2sm 1.07 1.93 4.87–8.79 21.95 1.46 33.86–46.34 16.17 17.31 S. leucophylla 26m + 4sm 0.95 1.81 5.01–9.55 18.96 1.26 35.00–43.28 17.62 20.83 S. mellifera 26m + 4sm 0.97 2.17 4.44–9.92 21.87 1.46 34.75–45.89 22.13 18.26 S. cardiophylla 0.82 1.61 3.70–7.26 22.19 1.23 S. cuspidata 1.05 1.88 3.43–6.14 30.63 1.39 S. splendens 28m + 16sm 0.72 1.38 3.23–6.20 22.26 1.01 27.27–45.79 18.23 23.67 S. subrotunda 2M + 26m + 16sm 0.75 1.42 3.18–6.01 23.62 1.07 25.96–50.00 15.32 21.92 S. microphylla 34m + 12sm 0.78 1.89 2.49–6.03 31.36 1.36 31.53–46.56 19.53 21.43 S. oppositiflora 0.53 1.21 2.31–5.27 22.98 0.82 S. stolonifera 48m + 12sm 0.81 1.73 2.19–4.69 36.92 1.23 25.49–46.34 18.79 22.40 S. atrocyanea 0.62 1.51 2.23–5.43 27.80 0.93 26 Halil Erhan Eroğlu et al. tion. Although Salvia is a polybasic genus with species of polyploid origin (Sheidai and Alijanpoo 2011), variations are observed resulting from dysploidy shows that differ- ent basic numbers with karyotypes that contain one or a few chromosomes more or less than that of the original, occur in a genus. S. microphylla has different basic num- ber (x = 23) probably with dysploidy. These data indicate that the effects of dysploidy on the lineage diversification of Salvia should be investigated further. In studied species, B-chromosomes, a special type of supernumerary chromosomes and are extra chromo- somes other than basic A-chromosomes in diploid and polyploid species, have been reported. The karyotype formulae are 22 + 0-1B in S. involucrata and 44 + 0-1B in S. cardiophylla and S. splendens (Alberto et al. 2003). We have not observed B-chromosomes. As a matter of fact, while B-chromosomes do not exist in some indi- viduals of the same population, the others may have dif- ferent numbers. When the number of B-chromosomes is small, they cannot have a visible effect on the phenotype and their presence can be determined only by cytologi- cal examinations. In case of high numbers, they have a negative effect on the development and fertility of plants (Houben 2017). A symmetric karyotype contains a high propor- tion of median and submedian chromosomes, unlike an asymmetric karyotype has a high rate of subterminal and terminal chromosomes (Peruzzi and Eroğlu 2013). In intrachromosomal asymmetry, the most symmetri- cal and asymmetrical karyotype are S. curviflora and S. corrugata, respectively. The relatively higher asymmetric karyotypes than other species may have been caused by chromosomal structural changes as centric fission or cen- tric fusion observed in especially polyploid and dysploidy species. In interchromosomal asymmetry, the most sym- metrical and asymmetrical karyotype are S. longifolia and S. mellifera, respectively. The relatively higher asym- metric karyotypes than other species may be the result of chromosome rearrangements and may also result in bimodality observed in S. campanulata, S. splendens, and S. microphylla. In these species, the bimodal karyotypes may occur due to loss of chromosomal segments follow- ing polyploidy. The symmetric and asymmetric karyo- types are different between intrachromosomal asym- metry and interchromosomal asymmetry with very low positive correlation (r = 0.157) (Figure 3). All studied Salvia species contain only median and submedian chro- mosomes and are symmetrical as a common condition Figure 2. Ideograms of Salvia: (A) S. corrugata, (B) S. campanulata, (C) S. curviflora, (D) S. elegans, (E) S. darcyi, (F) S. greggii, (G) S. invo- lucrata, (H) S. longifolia, (I) S. vitifolia, (J) S. mexicana, (K) S. apiana, (L) S. leucophylla, (M) S. mellifera, (N) S. cardiophylla, (O) S. cuspi- data, (P) S. splendens, (R) S. subrotunda, (S) S. microphylla, (T) S. oppositiflora, (U) S. stolonifera, (V) S. atrocyanea (scale bars: 1 μm). 27The new chromosomal data and karyotypic variations in genus Salvia in genus Salvia (Sheidai and Alijanpoo 2011; Doğan et al. 2019). On the contrary, Hu et al. (2016) reported that S. bulleyana Diels, S. digitaloides Diels and S. przewalskii Maxim. had asymmetrical karyotypes. In this study, it was aimed to determine the chro- mosome number of 21 Salvia species, to determine chromosome morphology, to reveal karyotype analy- sis in detail and to contribute to the cytotaxonomy of Salvia. In this context, the results are as follows: (i) the first report for the number of chromosomes of ten spe- cies, (ii) the karyotypic variations and new chromosome numbers different from previous reports for three spe- cies, (iii) the detailed chromosome measurements and karyotype analyses for all species studied for the first time, (iv) the symmetrical karyotypes for all studied species, (v) the variations resulting from dysploidy or polyploidy and discussing their reasons. On the other hand, the genus Salvia is one of the largest in the world with about 1000 species. The results of such studies pro- vide important data supports for Salvia cytotaxonomy. It is an important issue that combining all supporting data with further comparative studies and integrating them into morphological data. ACKNOWLEDGMENTS This work was supported by the [Uşak University Scientific Research Projects Fund] under Grant [number 2013/MF003]. REFERENCES Afzal-Rafii Z. 1976. 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