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Fluorescent Band Pattern of Chromosomes in Pseudolarix Amabilis, Pinaceae

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Fluorescent Band Pattern of Chromosomes in Pseudolarix Amabilis, Pinaceae © 2015 The Japan Mendel Society Cytologia 80(2): 151–157 Fluorescent Band Pattern of Chromosomes in Pseudolarix amabilis, Pinaceae Masahiro Hizume* Faculty of Education, Ehime University, 3 Bunkyo-cho, Matsuyama, Ehime 790–8577, Japan Received October 27, 2014; accepted November 18, 2014 Summary Pseudolarix amabilis belongs to one of three monotypic genera in Pinaceae. This species had 2n=44 chromosomes in somatic cells and its karyotype was composed of four long submetacentric chromosomes and 40 short telocentric chromosomes that varied gradually in length, supporting previous reports by conventional staining. The chromosomes were stained sequentially with the fluorochromes, chromomycin A3 (CMA) and 4′,6-diamidino-2-phenylindole (DAPI). CMA- bands appeared on 12 chromosomes at near terminal region and proximal region. DAPI-bands appeared at centromeric terminal regions of all 40 telocentric chromosomes. The fluorescent-banded karyotype of this species was compared with those of other Pinaceae genera considering taxonomical treatment and molecular phylogenetic analyses reported. On the basis of the fluorescent-banded karyotype, the relationship between Pseudolarix amabilis and other Pinaceae genera was discussed. Key words Chromomycin, Chromosome, DAPI, Fluorescent banding, Pinaceae, Pseudolarix amabilis. In Pinaceae, 11 genera with about 220 species are distinguished and grow mostly in the Northern Hemisphere (Farjon 1990). Most genera are evergreen trees, and only Larix and Pseudolarix are deciduous. Pinus is the largest genus in species number, and Cathaya, Nothotsuga and Pseudolarix are monotypic genera. The taxonomy of Pinaceae with 11 genera is complicated, having some problems in species or variety level. Several higher taxonomic treatments were reported on the base of anatomy and morphology such as resin canal in the vascular cylinder, seed scale, position of mature cones, male strobili in clusters from a single bud, and molecular characters in base sequences of several DNA regions. The family Pinaceae was divided into three subfamilies; Pinoideae (Pinus), Lariciodeae (Cedrus, Larix, Pseudolarix) and Abietoideae (Abies, Cathaya, Keteleeria, Picea Psudotsuga, Tsuga) by Pilger (1926), into two groups of Abies-Cedrus- Keteleeria-Pseudolarix-Tsuga and Cathaya-Larix-Picea-Pinus-Pseudotsuga by Melchior and Werdermann (1954), and into four subfamilies: Pinoideae (Pinus), Piceoideae (Picea), Laricoideae (Larix, Cathaya, Pseudotsuga), and Abietoideae (Abies, Cedrus, Pseudolarix, Keteleeria, Nothotsuga, Tsuga) by Hart (1987) and Frankis (1989). These classifications should be referred to Farjon (1990). Early molecular phylogenetic studies using PCR-PFLP and base sequence of rbcL (Tsumura et al. 1995, Chaw et al. 1997) indicated two groups in the phylogenetic tree but the trees had low reliability. Wu and Hu (1997) and Wang et al. (2000) presented three groups discussing relationships between or among genera on the basis of various morphology and anatomical structures. Recently, Gernandt et al. (2008) used a matrix of morphology, anatomical characters and base sequences of matK and rbcL and constructed reliable phylogenetic trees, which indicated that Pinaceae is divided two subfamilies, Abietoideae (Cedrus, Abies, Keteleeria, Pseudolarix, Nothotsuga, Tsuga) and Pinoideae (Larix, Psudotsuga, Cathaya, Picea, Pinus). The molecular * Corresponding author, e-mail: [email protected] DOI: 10.1508/cytologia.80.151 152 M. Hizume Cytologia 80(2) phylogenetic tree suggests that Pseudolarix is put in subfamily Abietoideae and relates to Tsuga and Nothotsuga. Most genera of Pinaceae are studied on basic chromosome number and their karyological information are deposited. Ten genera have common basic chromosome number, x=12 except for the monotypic genus, Pseudolarix possessing n=22 and 2n=44 and one species of Pseudotsuga, Pt. menziesii having n=13 and 2n=26 (Sax and Sax 1933, Khoshoo 1959, Mehra 1968, Hizume 1988). Karyotypes of species are common in each genus with variation in number or location of secondary constriction and slight change of chromosome shape. Karyotypes of the genera in Pinaceae were divided into several groups (Hizume 1988, Li 1995). Nkonggolo and Mehes-Smith (2012) reviewed recent molecular cytogenetic studies on Pinaceae karyotype implicating with molecular phylogeny. In Pinaceae most chromosomes are meta- and submeta-centric. Only two species, Pt. menziesii and Pl. amabilis, have telocentric chromosomes in their chromosome complements. Although five other Pseudotsuga species have 2n=24 chromosome number in common with Pinaceae genera and their karyotypes composed of 12 long metacentric chromosomes and 12 short submetacentric chromosomes, Pt. menziesii has 2n=26 chromosomes composed of 10 long metacentric chromosomes, 12 short submetacentrics and four short telocentrics. Barner and Christiansen (1962), Thomas and Ching (1968), El-Kassaby et al. (1983) and Sziklai et al. (1987) speculated that four telocentric chromosomes were derived from two pairs of long metacentric chromosomes by centromeric fission. Until now the explanation is not demonstrated cytogenetically by making a hybrid between 2n=26 and 2n=24 species and the observation of the meiotic configuration in the pollen mother cells of the hybrid. The karyotype (2n=44) of Pl. amabilis is composed of two pairs of submetacentric chromosomes and 20 pairs of telocentric chromosomes, which was explained with polyploidy and chromosome reduction (Sax and Sax 1933) and centromeric fission in the karyotype of certain Pinaceae genus (Khoshoo 1959, Mergen 1961, Gustafsson and Mergen 1964, Li 1994). Recently Zonneveld (2012) reported genome seizes in 172 conifer species including five genera of subfamily Abietoideae, 32–52 pg/C in Abies, 31–41 pg/C in Tsuga, 48.4 pg/C in Keteleeria, 32–40 pg/C in Cedrus, and 52.2 pg/C in Pseudolarix. The deviation of genome sizes suggests that Pl. amabilis might be diploid species. This study reports fluorescent banding pattern of chromosomes in Pl. amabilis and discusses its origin and phylogenetic position in Pinaceae. Materials and methods Small trees of Pseudolarix amabilis (Nelson) Rehder were obtained from commercial sources and planted in a pot. In the spring, growing root-tips were collected and treated in 2 mM 8-hydroxiquinoline or 0.05% colchicine for 6–7 h. Then the root-tips were immersed in a fixative (ethanol : acetic acid : chloroform=2 : 1 : 1) and stored in a freezer. Fixed root-tips were through 70% ethanol and immersed in water. Then the root-tips were soaked in 45% acetic acid for 5 min and transferred into 45% acetic acid at 60°C for 10 min. The root-tips were transferred into cold 45% acetic acid. Under a stereomicroscope, each root-tip was dissected by tweezers and a needle, and then the meristematic tissue was scooped and put on a glass slide. An aliquot of 45% acetic acid was dropped on the tissue and a cover glass was put on it. Cells of meristems were spread and squashed. The preparation was put on dry ice for a few minutes and the cover glass was ripped off. The preparation was air-dried overnight. Sequential fluorescent banding method with guanine specific CMA and adenine-thymine specific DAPI was described earlier by Kondo and Hizume (1982). The dried preparation was immersed into McIlvaine buffer pH 7.0 for 30 min. The glass slide was treated with 0.1 mg/mL distamycin A for 10 min, then washed with the buffer containing 5 mM MgSO4 for 10 min and stained with 0.1 mg/mL CMA in the buffer. After a wash with the buffer for 10 min the preparation was mounted with non-fluorescence glycerin and stored in a 2015 Fluorescent Band Pattern of Chromosomes in Pseudolarix amabilis, Pinaceae 153 refrigerator at 4°C for more that one night. After storage the CMA-stained preparation was observed under an epifluorescence microscope using a B filter cassette. Then the preparation was dipped in distilled water until the cover glass dropped off. Then the preparations were treated with acetic–alcohol (3 : 1) to remove CMA and glycerin, rinsed briefly with distilled water and then air- dried. The preparation was put in the buffer without MgSO4 for 10 min, treated with 0.1 mg/mL actinomycin D for 10 min and then washed again for 10 min with the buffer. The preparation was stained with 0.1 μg/mL DAPI for 5 min then washed with the buffer for 5 min. After mounting with the buffer–glycerin (1 : 1, v/v) mixture the same chromosomes observed with CMA were observed under the fluorescence microscope using a UV filter cassette. Fluorescence photographs of the same chromosomes stained with CMA and DAPI were taken on a film (TMAX, Kodak) and developed with double dilution of D-76. Results and discussion The chromosome number in somatic cells was observed to be 2n=44 in all six plants of Pl. amabilis examined. The chromosome complement was composed of four long submetacentric chromosomes and 40 short telocentric chromosomes, gradually decreasing in their lengths (Fig. 1). The somatic chromosome number and the karyotype observed were similar to previous studies (Sax and Sax 1933, Mergen 1961, Hizume 1988, Li 1994). After CMA-banding, 12 CMA-bands appeared in the somatic chromosome complement and most CMA-bands appeared at near region of centromeric ends (Fig. 1A). Ten CMA-bands were located at the distal position of DAPI-bands at the terminal or centromeric region, but not the terminal end of telocentric chromosomes. Two long submetacentric chromosomes had a somewhat weak CMA-band at the proximal region of their long arm. DAPI-bands appeared in all 40 telocentric chromosomes and were located at the centromeric or terminal region of telocentric chromosomes. Four long submetacentric chromosomes did not have any DAPI-bands (Fig. 1B). When the CMA-banding pattern was compared with the DAPI-banding pattern, the CMA-band was DAPI-negative and the DAPI-band was CMA-negative, same in other Pinaceae species and other Fig. 1. Fluorescent-banded somatic chromosomes of Pseudolarix amabilis at metaphase. (A) CMA staining. (B) DAPI staining. Bar=5 μm. 154 M. Hizume Cytologia 80(2) plant species. The karyotype and fluorescent banding pattern of Pl.
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