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Exploration of Intraspecific Cytomorphological Diversity in Agrimonia Eupatoria L

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Exploration of Intraspecific Cytomorphological Diversity in Agrimonia Eupatoria L © 2011 The Japan Mendel Society Cytologia 76(1): 81–88, 2011 Exploration of Intraspecific Cytomorphological Diversity in Agrimonia eupatoria L. (Rosaceae) from Western Himalayas, India Sanjeev Kumar, Syed Mudassir Jeelani*, Savita Rani, Santosh Kumari and Raghbir Chand Gupta Department of Botany, Punjabi University, Patiala 147 002, Punjab, India Received October 10, 2010; accepted January 18, 2011 Summary Agrimonia eupatoria L., a medicinally important species of the family Rosaceae, has been presently worked out for the first time from the 3 geographical areas of Kashmir (Jammu and Kashmir) and the Kangra and Sirmaur districts (Himachal Pradesh) of the Western Himalayas in India. The cytotypes with nϭ14 and nϭ28 were in conformity with the previous reports of the species from different parts of the world. A new cytotype of nϭ42 from Kashmir was observed for the first time. In comparison, these cytotypes (nϭ14, 28, 42) show significant variations in relation to morphology as well as geographical distribution in the Western Himalayas. Further, intra-population variability has been observed in different accessions of hexaploid cytotype in the form of B- chromosomes, abnormal meiotic course or aberrant type of flower morphology. Key words Agrimonia eupatoria, Cytotypes, Meiotic abnormalities, Morphovariants, Polyploidy, Western Himalayas. The genus Agrimonia L. typifies the tribe Sanguisorbeae of the sub-family Rosoideae in Rosaceae. The genus is herbaceous and composed of 15 species distributed in Northern and Southern temperate regions of the world (Santapau and Henry 1973). In India, the genus is represented by 2 species as A. eupatoria and A. pilosa (Hooker 1879). A. eupatoria L. is popularly known as “Agrimony” with 2 synonyms as A. pilosa var. neplensis (D. Don) Nakai and A. odorata (Gouan) Mill. and are distributed throughout the Himalayan range (1,800 to 3,500 m). They grow on open slopes as well as undergrowth in forests. The species is an erect, branched, smooth/hairy herb with pinnately compound leaves. Flowers are yellow in terminal spike-like racemes and the fruit is an achene with a ring of hooked bristles/spines above. The species are medicinally reputed with multiple uses such as anti-inhibitory activity against the hepatitis B virus (Dur et al. 2005) and as an anti-diabetic (Lakashman and Suneetha 2010). They are used in folk medicine as haemostatic, tonic for asthenia, astringent in diarrhoea and diuretic agents (Bilia et al. 1993). Materials and methods Morphological study In total, 24 morphometric characteristics (see Table 2) were studied for each cytotype to have proper insight of morphological variation. For stomata studies, mature leaves were treated with 10% aqueous solution of potassium hydroxide (KOH) at room temperature for 10–15 min and then epidermal peels so obtained were stained with 10% saffarnin in 95% ethanol. * Corresponding author, e-mail: [email protected] 82 S. Kumar et al. Cytologia 76(1) Table 1. Information about area, locality, altitude, accession number, meiotic chromosome number and ploidy level of different cytotypes of A. eupatoria from Western Himalayas Acession Meiotic Locality, Ploidy Meiotic S. No. number chromosome altitude (m) level course (PUP) number (n) Kangra District (Himachal Pradesh) 1. Dharmshala; 1,800 52545 14 2x Normal 2. Dharmkot; 2,000 53506 14 2x Normal 3. Palampur; 2,000 53511 28 4x Normal 4. Dharmshala; 2,100 53512 28 4x Normal 5. Baragran; 2,800 53507 28 4x Normal 6. Dal-Lake; 2,800 53510 28 4x Normal 7. Lohardhari; 3,000 53508 28 4x Normal 8. Rajgundha; 3,000 53509 28 4x Normal Sirmaur District (Himachal Pradesh) 9. Timbi; 1,800 53498 14 2x Normal 10. Dedag; 1,800 53494 14 2x Normal 11. Nauradhar; 1,900 52578 28 4x Normal 12. Haripur dhar; 2,600 52571 28 4x Normal 13. Chur Dhar; 2,300 53492 28 4x Normal 14. Bhangyani mata; 2,800 53493 28 4x Normal 15. Tisri; 3,000 53496 28 4x Normal Kashmir (Jammu and Kashmir) 16. Aharbal; 2,100 53478 28 4x Normal 17. Keller; 2,300 52487 28 4x Normal 18. Yusmarg; 2,400 53475 28 4x Normal 19. Hirpora; 2,400 53480 28 4x Normal 20. Sonmarg; 3,000 53481 42 6x Abnormal 21. Lidderwatt; 3,100 53474 42 6x Abnormal 22. Chumnai; 3,500 53476 42ϩ (0-1B) 6x Normal 23. Aharbal; 3,200 52488 42 6x Normal 24. Chumnai, 3,500 53482 42 6x Abnormal Cytological studies For meiotic studies, flower buds were collected from selected localities of diverse different areas of Western Himalayas (Table 1). Smears of appropriate sized flower buds were made after fixing these in Carnoy’s fixative, using the standard acetocarmine technique. Pollen fertility was estimated by mounting mature pollen grains in glycero–acetocarmine (1 : 1) mixture. Well-filled pollen grains with stained nuclei were taken as apparently fertile, while shrivelled and unstained pollen grains were counted as sterile. Photomicrographs of pollen mother cells and pollen grains were made from freshly prepared slides using Nikon 80i eclipse Digital Imaging System. Voucher specimens are deposited in the Herbarium, Department of Botany, Punjabi University, Patiala (PUN). Results Morphological observations For the morphological studies, 24 parameters have been evaluated for vegetative and reproductive characteristics of all 3 cytotypes (Table 2). Significant morphological differences are noted for the characteristics such as plant height, leaves (number, size of the leaflets and, stomata size and index), length of spike of inflorescence and shape of bristles of fruit. In one of the accessions (52488) of hexaploid cytotype, 6 sepals and petals were reported instead of the normal 5. 2011 Exploration of Intraspecific Cytomorphological Diversity in Himalayan Rosaceae 83 Table 2. Detailed morphological comparison of 3 cytotypes of A. eupatoria from different areas of Western Himalayas Morphological Diploid Tetraploid Hexaploid No. characteristics (nϭ14) (nϭ28) (nϭ42) 1. Distribution Uncommon Most Common Uncommon 2. Habit Small sized herb Medium sized herb Tallest 3. Habitat Found under Found along roadsides Found on the slopes shade of in on humus bushes in forests rich places in forests 4. Plant height (cm) 30–40 60–90 70–120 Stem 5. Surface Non-glabrous Non-glabrous Glabrous 6. No. of hairs/mm2 — — 10–50 7. Length of hairs (cm) — — 0.2–0.5 Leaf 8. Number of leaves/plant 6–8 10–12 8–13 Leaflets 9. Shape Ovate Lanceolate Lanceolate 10. Average size (cm) 2.1ϫ1.1 3.1ϫ1.6 6.8ϫ1.8 11. Number 4–6 6–8 5–7 12. Apex blunt blunt pointed Stipule 13. Length (cm) 0.2–0.8 0.4–1.2 0.2–1.4 Stomata 14. Size (mm) 23.65ϫ18.92 24.83ϫ19.51 27.40ϫ20.41 15. Stomatal frequency on upper/ 5.46/3.24 5.79/3.48 6.25/3.25 lower surface of leaf (mm2) 16. Stomatal index of upper/ 22.28/12.45 23.45/13.24 24.50/14.32 Inflorescence lower surface of leaf (mm) 17. Length of spike (cm) 8–12 15–19 18–28 Flower 18. Size (cm) 0.4ϫ0.3 0.4ϫ0.3 0.5ϫ0.3 19. Number of stamens/flower 7–10 7–10 8–15 Fruit 20. Size (cm) 0.3–0.5 0.3–0.5 0.5–0.7 21. Shape of bristles Slightly curved Curved More curved 22. Number of achenes/fruit 1 1 (2 in Acc. No. 52571) 1 Pollen grains 23. Size (mm) 22.53ϫ19.54 28.15ϫ24.16 28.52ϫ22.19–35.15ϫ33.50 24. Shape Round Oblong Round-Quadrangular Cytological observations Based on xϭ14, the 24 different accessions of A. eupatoria investigated at present from the Western Himalayas revealed the existence of diploid (2nϭ28), tetraploid (2nϭ56) and hexaploid (2nϭ84) cytotypes. Two accessions from Kangra and 2 from Sirmaur were diploids (nϭ14) showing 14II at metaphase-I (M-I, Fig. 1). Six accessions from Kangra, 5 from Sirmaur and 4 from ϭ Kashmir were tetraploids (n 28) showing 28II at diakinesis (Fig. 2). The other 5 accessions from ϭ Kashmir were hexaploids (n 42) showing 42II at diakinesis (Fig. 3) with accession number 53476 depicting 0-1B chromosome per PMC (Fig. 4). The course of meiosis and microsporogenesis in these accessions was observed to be normal with high pollen fertility except for the 3 accessions of hexaploid cytotype exhibiting abnormal meiosis and reduced pollen fertility as shown in Table 3. The various meiotic abnormalities in these accessions being observed were unoriented bivalents at M-I (Fig. 5), chromosomal bridges and laggards at anaphases and telophases (Figs. 6, 7) and 84 S. Kumar et al. Cytologia 76(1) Figs. 1–20. 1–7. Photomicrographs of PMCs of A. eupatoria. 1. Diakinesis-I (nϭ14), 2. Diakinesis ϭ ϭ ϩ ϭ ϩ (n 28), 3. Diakinesis (n 38II 2IV), 4. M-I (n 42 1B), 5. M-I with unoriented bivalents, 6. A-I showing chromosomal bridge, 7. Telophase-II showing chromosomal laggards, 8. Triad with micronuclei. 9–12. Variable shape of pollen grains: 9. Round in diploid, 10. Oblong in tetraploid, 11. Round in hexaploid 12. Quadrangular in accession 52488 of hexaploid. 13–16. Stomatal size variations. 13. Diploid, 14. Tetraploid, 15. Hexaaploid, 16. in accession 52488 hexaploid. 17–20. Fruit surface bristle shape variation, 17. Slightly curved in diploid, 18. Curved in tetraploid, 19. More curved in hexaploid, 20. Extremely curved in accession 52488 of hexaploid. Scaleϭ10 mm. formation of micronuclei (Fig. 8). A significant difference is seen in the size and shape of pollen grains in these cytotypes (Figs. 9–11). Geographical distribution In the Western Himalayas, the distribution pattern of euploid cytotypes showed a definite relation to altitudinal variations (Table 1). The diploids in the Himachal Pradesh were uncommon but some accessions were available from lower altitudes of 1,800–2,000 m.
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