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Cytogeography of Glechoma Hederacea Subsp. Grandis (Labiatae) in Japan

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© 2010 The Japan Mendel Society Cytologia 75(3): 255–260, 2010 Cytogeography of Glechoma hederacea subsp. grandis (Labiatae) in Japan Norihito Miura and Yoshikane Iwatsubo* Graduate School of Science and Engineering, University of Toyama, Gofuku 3190, Toyama 930–8555, Japan Received February 26, 2010; accepted August 28, 2010 Summary In this study, we examined the chromosomal number for Glechoma hederacea subsp. grandis in a total of 1,030 specimens collected from different distribution areas in Japan. We found that G. hederacea subsp. grandis could be categorized into 3 cytotypes with 2nϭ36 (tetraploid), 2nϭ45 (pentaploid) and 2nϭ54 (hexaploid) chromosomes. Tetraploid plants were found throughout different areas in Japan; however, hexaploid plants were mainly distributed in central Honshu, Shikoku and Kyushu. Likewise, pentaploid plant distribution was found to overlap with hexaploid plant distribution areas. The pentaploid plant group appeared only in regions common to both tetraploid and hexaploid plants. All 3 cytotypes were found to have karyotypes which could be represented by the following equations: A) 6Mϩ4mϩ18smϩ8st for tetraploids, B) 6Mϩ15mϩ19smϩ5st for pentaploids, and C) 6Mϩ26mϩ20smϩ2st for hexaploids. Pentaploid specimen karyotypes had half the tetraploid and half the hexaploid chromosomal set, indicating that this specimen was a hybrid between tetraploid and hexaploid plants. Key words Geographic distribution, Glechoma hederacea subsp. grandis, Hybrid, Karyotype, Polyploidy. Glechoma L. (Labiatae), distributed across north temperate zones in Eurasia, is a small genus with 4 to 8 species (Budantsev 2004). One of its species, G. hederacea L., has a wide distribution range occurring spontaneously throughout Eurasia. Furthermore, this species can be divided into subsp. hederacea distributed in Europe, and subsp. grandis (A. Gray) H. Hara distributed in Far East Asia (Hara et al. 1954). The subsp. grandis (Fig. 1) is the only taxon of this genus found in Japan and known to have a chromosomal count of 2nϭ36 (Tanaka 1953, Hara et al. 1954, Nishikawa 1985). However, Iwatsubo et al. (2004) revealed that G. hederacea subsp. grandis located in Toyama Prefecture, situated in the central region of Honshu in Japan, had 3 cytotypes corresponding to 2nϭ36, 45 and 54, and interpreted these as tetraploids, pentaploids, and hexaploids, respectively. Moreover, the authors concluded Fig. 1. Photograph of plant of Glechoma hederacea that of the 3 chromosomal forms, the pentaploid subsp. grandis. * Corresponding author, e-mail: [email protected] 256 N. Miura and Y. Iwatsubo Cytologia 75(3) form was a hybrid between tetraploid and hexaploid plants (Iwatsubo et al. 2004). A recent study by Miura and Iwatsubo (2008) found that G. hederacea subsp. grandis was also distributed in Miyagi Prefecture, the Tohoku districts of Honshu. Their study also revealed that most plants were tetraploids, with only a fraction of hexaploid plants. The studies conducted from the 2 prefectures indicated that each chromosomal form of G. hederacea subsp. grandis could have different distributions in Japan. Therefore, we decided to study the distribution and karyotype of the 3 cytotypes of this subspecies in order to shed light on the putative origin of the pentaploid plant. Materials and methods A total of 1,030 specimens of G. hederacea L. subsp. grandis were collected from 945 localities in Japan. Newly formed adventitious root tips were collected and pretreated with 2 mM 8- hydroxyquinoline solution at room temperature for 1–1.5 h, and maintained at approximately 5°C for 15 h. The root tips were then fixed with a mixture of glacial acetic acid and absolute ethyl alcohol (1 : 3), and incubated at room temperature for 1 h. The root tips were then macerated in 1 N hydrochloric acid at 60°C for 10 min, and washed in tap water. Finally, the samples were stained and squashed in a drop of 1.5% lacto-propionic orcein on a glass slide, and fully-spread metaphase chromosomes were examined. Collection localities for the specimens were as follows: A) tetraploid plants from Higashikuromaki, Toyama City, Toyama Prefecture, B) pentaploid plants from Hio, Toyama City, Toyama Prefecture, and C) hexaploid plants from Monjuji, Toyama City, Toyama Prefecture. Chromosomal forms were expressed according to the nomenclature outlined by Levan et al. (1964). Voucher specimens of the plants examined have been deposited at the Toyama Science Museum (TOYA). Results and discussion Chromosome numbers Chromosomal counts determined in this study for G. hederacea subsp. grandis were based on 1,030 specimens collected from the distribution areas of Japan known to have the 3 cytotypes (2nϭ36: tetraploids, 2nϭ45: pentaploids, 2nϭ54: hexaploids) of this taxon (Fig. 2). These distribution areas almost corresponded to previous reports for this taxon in Japan (Tanaka 1953, Hara et al. 1954, Nishikawa 1985, Iwatsubo et al. 2004, Miura and Iwatsubo 2008). From a total of 1,030 specimens, 817 (79.3% of the plants examined) were tetraploids with 2nϭ36 chromosomes, 83 (8.1%) were pentaploids with 2nϭ45 and 130 (12.6%) were hexaploids with 2nϭ54. Fig. 2. Photographs of somatic metaphase chromosomes of 3 chromosome forms of Glechoma hederacea subsp. grandis, A: 2nϭ36; B: 2nϭ45; C: 2nϭ54. Bar indicates 5 mm. 2010 Cytogeography of Glechoma hederacea subsp. grandis (Labiatae) in Japan 257 Fig. 3. Distribution of area of the 3 chromosome forms of Glechoma hederacea subsp. grandis in Japan, A: 2nϭ36; B: 2nϭ45; C: 2nϭ54. Bar indicates 200 km. Fig. 4. Karyotypes of somatic metaphase of 3 chromosome forms of Glechoma hederacea subsp. grandis, A: 2nϭ36; B: 2nϭ45; C: 2nϭ54. Bar indicates 5 mm. 258 N. Miura and Y. Iwatsubo Cytologia 75(3) Distribution areas for the 3 cytotypes were as follows: A) Tetraploids (Fig. 3A) occurred throughout Hokkaido, Honshu, Shikoku, and Kyushu. In Hokkaido, all plants were tetraploids. Almost all plants located in the northern part of Honshu (Tohoku and Kanto districts) were also tetraploid. B) Pentaploids (Fig. 3B) were distributed in central Honshu, Shikoku and Kyushu. However, this plant occurred only in the distribution area common to the hexaploid plant. C) Hexaploids occurred mainly in central Honshu, Shikoku and Kyushu (Fig. 3C). Karyotype analysis Table 2. Masurements of somatic metaphase chromosomes of 2nϭ45 Glechoma hederacea Tetraploid plants had chromosomes at subsp. grandis metaphase which ranged from 0.9 mm to Chromosome 2.2 mm in length, and had an arm ratio between No. Length (mm) Arm ratio 1.0 and 4.5 (Fig. 4A). As shown in Table 1, the form 36 chromosomes were divided into 3 groups: 1 0.3ϩ1.4ϭ1.7 4.7 st 2 0.3ϩ1.3ϭ1.6 4.3 st Table 1. Measurements of somatic metaphase chromo- 3 0.4ϩ1.2ϭ1.6 3.0 sm somes of 2nϭ36 Glechoma hederacea subsp. 4 0.4ϩ1.2ϭ1.6 3.0 sm grandis 5 0.4ϩ1.2ϭ1.6 3.0 sm 6 0.5ϩ1.1ϭ1.6 2.2 sm Chromosome 7 0.5ϩ1.1ϭ1.6 2.2 sm No. Length (mm) Arm ratio form 8 0.6ϩ1.0ϭ1.6 1.7 m 9 0.6ϩ0.9ϭ1.5 1.5 m 1 0.4ϩ1.8ϭ2.2 4.5 st 10 0.6ϩ0.9ϭ1.5 1.5 m 2 0.4ϩ1.7ϭ2.1 4.3 st 11 0.3ϩ1.2ϭ1.5 4.0 st 3 0.4ϩ1.5ϭ1.9 3.8 st 12 0.3ϩ1.2ϭ1.5 4.0 st 4 0.4ϩ1.5ϭ1.9 3.8 st 13 0.3ϩ1.1ϭ1.4 3.7 st 5 0.4ϩ1.4ϭ1.8 3.5 st 14 0.4ϩ1.0ϭ1.4 2.5 sm 6 0.4ϩ1.4ϭ1.8 3.5 st 15 0.4ϩ1.0ϭ1.4 2.5 sm 7 0.4ϩ1.4ϭ1.8 3.5 st 16 0.4ϩ1.0ϭ1.4 2.5 sm 8 0.4ϩ1.4ϭ1.8 3.5 st 17 0.5ϩ0.9ϭ1.4 1.8 sm 9 0.5ϩ1.3ϭ1.8 2.6 sm 18 0.5ϩ0.8ϭ1.3 1.6 m 10 0.5ϩ1.3ϭ1.8 2.6 sm 19 0.6ϩ0.7ϭ1.3 1.2 m 11 0.5ϩ1.2ϭ1.7 2.4 sm 20 0.6ϩ0.7ϭ1.3 1.2 m 12 0.5ϩ1.2ϭ1.7 2.4 sm 21 0.3ϩ0.9ϭ1.2 3.0 sm 13 0.4ϩ1.2ϭ1.6 3.0 sm 22 0.3ϩ0.9ϭ1.2 3.0 sm 14 0.4ϩ1.2ϭ1.6 3.0 sm 23 0.4ϩ0.8ϭ1.2 2.0 sm 15 0.5ϩ0.9ϭ1.4 1.8 sm 24 0.4ϩ0.8ϭ1.2 2.0 sm 16 0.5ϩ0.9ϭ1.4 1.8 sm 25 0.4ϩ0.8ϭ1.2 2.0 sm 17 0.7ϩ0.7ϭ1.4 1.0 M 26 0.4ϩ0.8ϭ1.2 2.0 sm 18 0.7ϩ0.7ϭ1.4 1.0 M 27 0.5ϩ0.7ϭ1.2 1.4 m 19 0.7ϩ0.7ϭ1.4 1.0 M 28 0.5ϩ0.7ϭ1.2 1.4 m 20 0.7ϩ0.7ϭ1.4 1.0 M 29 0.5ϩ0.7ϭ1.2 1.4 m 21 0.5ϩ0.8ϭ1.3 1.6 m 30 0.5ϩ0.7ϭ1.2 1.4 m 22 0.5ϩ0.8ϭ1.3 1.6 m 31 0.5ϩ0.6ϭ1.1 1.2 m 23 0.4ϩ0.9ϭ1.3 2.3 sm 32 0.4ϩ0.7ϭ1.1 1.8 sm 24 0.4ϩ0.9ϭ1.3 2.3 sm 33 0.4ϩ0.7ϭ1.1 1.8 sm 25 0.6ϩ0.7ϭ1.3 1.2 m 34 0.3ϩ0.7ϭ1.0 2.3 sm 26 0.6ϩ0.7ϭ1.3 1.2 m 35 0.3ϩ0.7ϭ1.0 2.3 sm 27 0.6ϩ0.6ϭ1.2 1.0 M 36 0.4ϩ0.6ϭ1.0 1.5 m 28 0.6ϩ0.6ϭ1.2 1.0 M 37 0.5ϩ0.5ϭ1.0 1.0 M 29 0.3ϩ0.8ϭ1.1 2.7 sm 38 0.5ϩ0.5ϭ1.0 1.0 M 30 0.3ϩ0.8ϭ1.1 2.7 sm 39 0.5ϩ0.5ϭ1.0 1.0 M 31 0.4ϩ0.7ϭ1.1 1.8 sm 40 0.5ϩ0.5ϭ1.0 1.0 M 32 0.4ϩ0.7ϭ1.1 1.8 sm 41 0.4ϩ0.5ϭ0.9 1.3 m 33 0.4ϩ0.7ϭ1.1 1.8 sm 42 0.4ϩ0.5ϭ0.9 1.3 m 34 0.4ϩ0.7ϭ1.1 1.8 sm 43 0.4ϩ0.4ϭ0.8 1.0 M 35 0.3ϩ0.7ϭ1.0 2.3 sm 44 0.4ϩ0.4ϭ0.8 1.0 M 36 0.3ϩ0.6ϭ0.9 2.0 sm 45 0.3ϩ0.4ϭ0.7 1.3 m 2010 Cytogeography of Glechoma hederacea subsp.
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  • Table S1. Sambles, Vouchers, and Accession Number of ETS and ITS Sequences Used in This Study. Accession Number in Bold Letter

    Table S1. Sambles, Vouchers, and Accession Number of ETS and ITS Sequences Used in This Study. Accession Number in Bold Letter

    Table S1. Sambles, vouchers, and accession number of ETS and ITS sequences used in this study. Accession number in bold letter indicates the sequences determined in this study. Taxon name code ETS ITS voucher References Agastache pallida (Lindl.) Cory JQ669144 JQ669075 B.Drew 118 Agastache rugosa Kuntze JQ669145 JQ669076 H.Kanai, K Hasegawa, K.Ohkubo 8916 Caryopteris incana Miq. KY552620 EF508064 PNLI20120421-1 Li et al. 2017 Glechoma longituba (Nakai) Kuprian. KM886687 KM886722 T.Deng 415 (KUN) Deng et al. 2015 Glechoma biondiana (Diels) C.Y.Wu & C.Chen KM886693 KM886728 D.G.Zhang 4446(KUN) Deng et al. 2015 Glechoma biondiana (Diels) C.Y.Wu & C.Chen KM886691 KM886726 D.G.Zhang 4731(KUN) Deng et al. 2015 Glechoma biondiana v. angustituba C.Y.Wu & C.Chen KM886685 KM886720 D.G.Zhang 4583(KUN) Deng et al. 2015 Glechoma hederacea L. JQ669171 JQ669099 B.Drew 69 Hyptis laniflora Benth, JF304259 JF301548 JP Lewis 2034(K) unpublished Lamium album L. JX893206 JX893229 Liu and Xiang 128 Lycopus cavaleriei H. Lév. China KM886695 KM886730 SNJ Exped.20110807071(KUN) Lycopus cavaleriei H. Lév. Japan LC202992 LC542952 J.Oda 8240(KYO) Takano & Oda 2017, present study Lycopus uniflorus Michx. LC202993 LC542953 J.Oda & A.Muranaga 8555(KYO) Takano & Oda 2017, present study Lepechinialeucophylloides (Ramamoorthy, Hiriart & JF301327 JF301354 B.Drew 129 Drew & Sytsma 2011. Syst.Bot. 36: 1038-1049 Medrano) B.T.Drew, Cacho & Sytsma Nepeta cataria L. JQ669202 JQ669126 B.Drew 72 Drew & Sytsma 2012 Melissa officinalis L. JF301325 JF301353 B.Drew 70 Drew & Sytsma 2011. Syst.Bot. 36: 1038-1049 Prunella vulgaris L.
  • Ethnobotanical Review of Wild Edible Plants of Slovakia

    Ethnobotanical Review of Wild Edible Plants of Slovakia

    Acta Societatis Botanicorum Poloniae Journal homepage: pbsociety.org.pl/journals/index.php/asbp INVITED REVIEW Received: 2012.01.15 Accepted: 2012.08.26 Published electronically: 2012.11.16 Acta Soc Bot Pol 81(4):245–255 DOI: 10.5586/asbp.2012.030 Ethnobotanical review of wild edible plants of Slovakia Łukasz Łuczaj* Department of Botany and Biotechnology of Economic Plants, University of Rzeszów, Werynia 502, 36-100 Kolbuszowa, Poland Abstract This paper is an ethnobotanical review of wild edible plants gathered for consumption from the 19th century to the present day, within the present borders of Slovakia. Twenty-four sources (mainly ethnographic) documenting the culinary use of wild plants were analysed. The use of 106 species (over 3% of the Slovak flora) has been recorded. Nowadays most of them are no longer used, or used rarely, apart from a few species of wild fruits. The most frequently used plants include the fruits of Rubus idaeus, Fragaria spp., Rubus subgenus Rubus, Vaccinium myrtillus, V. vitis-idaea, Fagus sylvatica, Corylus avellana, Prunus spinosa, Pyrus spp., Malus spp., Crataegus spp. and the leaves of Urtica dioica, Rumex acetosa, Chenopodiaceae species, Cardamine amara, Glechoma spp., Taraxacum spp. and Oxalis acetosella. The most commonly used wild food taxa are nearly identical to those used in Poland, and the same negative association of wild vegetables with famine exists in Slovakia, resulting in their near complete disappearance from the present-day diet. Keywords: historical ethnobotany, ethnobiology, wild green vegetables, wild food plants, wild edible plants Introduction of rural populations started at the end of the 19th century.