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Reproductive Biology of Picrorhiza kurroa – a Critically Endangered High Value Temperate Medicinal Plant
R. Raina*, Tara Singh Mehra, Romesh Chand and Yash Pal Sharma
Department of Forest Products, Dr. Y.S. Parmar University of Horticulture and Forestry, Nauni- Solan (HP) 173 230 *Corresponding author: firstname.lastname@example.org
Abstract: Picrorhiza kurroa Royle ex. Benth.(Family: Scrophulariaceae), an important endangered temperate medicinal plant used as hepatic stimulant is a perennial herb propagated mostly by stolon segments. Chromosome number and its reproductive behaviour are reported for the first time. Meiosis is normal in the species with 2n=34 chromosomes. P. kurroa mostly prefers out-crossing as evinced by pollination experiments and floral architecture. Key Words: Picrorhiza kurroa, endangered medicinal plant, reproductive biology Manuscript received: 5 February, 2010 Manuscript accepted: 3 June, 2010 Methods Male meiotic studies were conducted by fixing unopened flower buds in Carnoy’s fixative II consisting of absolute alcohol: glacial acetic acid: chloroform (1:1:1) for 24 hours. One percent propionocarmine was used for staining chromosomes by the usual squash method and a minimum of 20 good cells at proper stage were studied for taking observations. Pollen stainability was determined by staining freshly dehisced pollens with 1% propionocarmine. Pollination studies were conducted with three treatments viz., (i) open pollination, (ii) selfing by bagging of single spikes and (iii) selfing 2-4 spikes/ plant with seven replications in randomized block design. Minimum 10 floral spikes were used in each replication.
Picrorhiza kurroa Royle ex. Benth.(Family: Scrophulariaceae), native to Western Himalayan region, between 3000-5000 m elevation (Hooker 1885; Chopra & Ghosh 1934; Blatter 1984; Jain 1996; Agrawal 2003) is valued as hepato-protective, antiperiodic, cholagouge, stomachic, anti-amoebic, anti-oxidant, anthelmintic, anti-inflammatory, cardio-tonic, laxative, carminative, expectorant, etc. (Chopra & Ghosh 1934; Uphof 1959; Kapoor 1990; Kapahi et al. 1993; Singh et al. 2006; Bhatt & Bhatt 1996: Gaddipati et al. 1999; Prajapati, 2003). Picroside I and II are the two important chemical constituents present in its rootstock which have therapeutical importance (Dutt et al. 2004). In order to fulfil the increasing national and international demand, the raw drug is largely wild extracted since the species has very limited cultivation that has made it critically endangered (Nayer and Sastry 1987; Rai et al. 2000). The species has also been put in the negative list of exports by Ministry of Commerce, GOI [Vide Notification No. 03 (RE2003)/2002-2007 (Appendix II) dated 31st March, 2003 issued by Director General of Foreign Trade, Govt. of India]. Despite considerable therapeutical importance, no information regarding its reproductive biology, ploidy status and breeding systems is available in literature. The present study was undertaken to understand its breeding behaviour that is fundamental to undertake any genetic improvement for higher yield and quality attributes.
Results and Discussion
Morphology Plants of P. kurroa are perennial, herbaceous with creeping rootstock and aerial parts represented by basal leaves and flowering scape only (Fig.1). Such features make P. kurroa highly adaptable to harsh environmental conditions of the areas it inhabits. The plants spread through stolons and apparently separate looking plants at above ground level are actually joined together by stolons beneath. However, when these stolons get detached from the mother stock, the plants become independent. The simple leaves arising from the tip of the upturned stolons are present in rosettes or whorls. Cauline leaves are present only during the flowering phase. The inflorescence is indeterminate terminal spike forming a more or less triangular head. The flowers are sessile, zygomorphic, bilipped, bisexual and purple (Fig. 2) borne in the axils of small green
Material and methods
Plant Material Wild and field grown plants of the species at Rahla, Himachla Pradesh, India (2800m amsl) were used for the study.
73 (2.56 (61. One can have a reasonable clue to the evolutionary primitiveness and/or advancement of a species based on certain morphological features (Cronquist 1968). the anthers are borne on incurved filaments. This ensures an efficient seed dispersal mechanism as the plants of this species inhabit high wind zones and the wind carries its seeds far away from the mother plant. This is probably to add colour to the flower for attracting insects for affecting pollination. introrse anthers ensure that some pollen may fall on the flowering spike itself so as to affect selfpollination. autogamy is often the only possible means of sexual reproduction (Hagerup 1932 & 1951). The anthers start shedding pollen once they come out of the corolla cavity till the maximum growth of the filament. 3 Propionocarmine stained pollen. as in high alpine or arctic region and deserts those are poor in pollinators because of the unfavourable conditions. Although exerted stamens are a step towards cross pollination. 1-9: Picrorhiza kurroa 1 Flowering plant. become purplish after anthesis. Although the stamens are exerted.00 (1.71) 1.setting parent. Bombus spp. 5 Seed germination in nursery.57 (4. 9 Metaphase-I (2n=34) 41 .05) Opened flowers/ spike (no.)** 17. which make their position introrse.29 21.50 mm) and 2 short (13.0 mm) and light seeds (1000 seeds weigh approximately 60 mg) (Fig.72) 0 (0. It also avoids overcrowding around the seed. The ovoid.84 (2.kurroa are long- 愀 戀 挀 搀 ㈀ ㌀ 㐀 㔀 㘀 㜀 㠀 㤀 Figs.117 (values in parenthesis are *arc sign transformed values. 2 Floral parts (a) single flower (b) calyx (c) corolla with four stamens (d) pistil.43 NS Fruits /spike (no. The flower architecture mostly favours cross-pollination. The four free stamens are epipetalous and alternating with the petal lobes. 8 Diakinesis(2n=34).933 Fruit size (mm)** 7. Sepals persist up to capsule development stage and are of unequal in size. In environments. 6 Effect of open and self pollination on spike length. However.71) 6.OAJMAP (2010) Table 1.25) 0. As is known (Kaul & Kaul 1996) and as well observed during the present studies.02 (17. which are colourless in bud stage. this species occurs at high altitudes characterized by short growing seasons. turgid and acute dry dehiscent capsules of this species dehisce septicidally and/ or loculicidally shedding extremely small sized (1. Treatments No. Effect of different pollination methods on fruit set in P. 3. The seed is covered by hyaline reticulate loose sac like seed coat enclosing the minute embryo.57 (1. **sq.61) 0. Selfing (by bagging of single inflorescence) Selfing (by caging of 2-4 inflorescences per plant) CD (P=0. Stamens are didynamous with 2 long (15.4).) 22.227 Fruit set (%)* 77.25 mm) stamens. in Picrorhiza kurroa. Simultaneous with anthesis the stamens become exerted with anthers continuing in the introrse position.5).49) 0. the primitive characters observed in P.14 21. Based on these (Cronquist 1968).43 (4.83) 0 (0) 9. Approximately 41% seeds germinated under laboratory conditions as compared to 35% under nursery conditions (Fig. and Apis florae were observed visiting the flowers. Open pollination 2. This was also observed in pollination studies (discussed later).46) 1. 1.95) 0 (0.17) 0 (0. root transformed values) coloured bracts opening during June to September. kurroa. 1. 4 Seeds.30 x.71) 2. Filaments. Sr. 7 Diplotene (2n=34). the exerted stamens as well as introrse anthers ensure both allogamy and autogamy. The stye also keeps pace with the filaments and at the stage of maximum length both were of almost equal length.101 Seed set/spike (mg)** 24.
Base numbers of above 12 are considered secondary base numbers having evolved as a result of ascending or descending dysploidy and a cross between forms with different base numbers followed by chromosome doubling (Stebbins 1971). approximately 9 per cent fruit set was observed when 2-4 spikes of the same plant were bagged together. simple chlorophyllous leaves. In none of the pollen mother cells could any abnormality like laggards. pp. It appears that as an insurance against failed crosspollination. www. where as temperate habitat. and Ghosh. Despite extensive literature survey. N. 3 species to Verbascum and 1 species to Veronicum. It also suggests that the species is cross pollinated and in the absence of the agents/ conditions. D. when more number of spikes of a single plant was bagged together. Indian Journal of Gastroenterology 15: 63-67. 17 bivalents were regularly observed at diplotene. petals united. etc be observed. Some common indigenous remedies.rothamsted-international. cyclic floral parts with few of each kind. S. no published record of cytogenetical details including the chromosome number of P. R. some extent of self pollination is also possible and self incompatibility barriers are not completely expressed. the introrse anthers ensure setting of some seed through self. Dutt.org/HTML/ Publications/Newsletters/DonorDec04. This view is also supported by floral morphology of this species (exerted stamens and free hanging stigma. A. and Bhatt. 1996.57 mg) under open pollination as compared to 1. Pollen stainability was found to be approximately 95%. the present observations of 2n=34 in P. 2003. 1984. tribe (Digitaleae) and series (Rhinanthideae) of family Scrophulariaceae. pollination by insects other than beetles (Bombus spp. carpel fully closed and united with axile placentation and terminal stigma.6). S. Due to the absence of any literature regarding chromosome number of the species. Kiddle. and Apis florae) are the major advanced characters. Cronquist. 2004. Indigenous drugs and liver disease. E. Seed yield per spike was also significantly higher (24. C..73 mm as compared to 6. The evolution and classification of flowering plants. hypogynous ovary forming dehiscent fruit and epigeal germination. One possibility could be of a cross between progenitors with x=8 and 9 followed by chromosome doubling. Development of fruit set. Pollination studies Approximately 77 per cent fruit set was observed in open pollinated spikes whereas no fruit set was observed when single spikes were bagged. kurroa appears to be the first report of chromosome number in this species. 8&9). B.pollination. 23 species belong to genus Veronica. diakinesis and metaphase-I followed by regular anaphase -I with chromosomes segregation of 17-17 univalents at each pole (Fig. This indicates that although the species favours cross-pollination. Khambay. London : Staples and Staples Ltd. H. V. The length of open pollinated spikes increased considerably whereas self pollinated ones did not show any length increase and subsequently dried with dry floral parts remaining attached (Fig. 42 . Differential accumulation of picrosides in Picrorhiza kurroa Royle ex Benth plants. A. 2 species to genus Celsia. Boston: Houghton Miffin.pdf. Cytological studies In all the pollen mother cells studied.. and Foyer. Dehradun: Bishen Singh Mahendra Pal Singh. micronuclei. S.57 mg recorded when 2-4 spikes/plant were bagged together (Table-1). Out of these 29 species. kurroa could be found. N. 143-145. References Agrawal.44 mm when 2-4 spikes/plant were bagged together. However. herbaceous and deciduous habit. indicates some degree of self pollination. Chopra. No fruit set under selfed (bagging) conditions and significant fruit set under open conditions were indications towards cross compatible behaviour of this species.OAJMAP (2010) lived habit. S. the presumed base number would be 17. Blatter. Indian Journal of Medical Research 22: 263-264. Singh.1968. stamens differentiated into filaments and anthers producing triaperturate pollen (Fig. The primary base numbers reported in family Scrophulariaceae form a series from x=6 up to 12 and taxa with 2n=12 to 168 are reported in literature (Federov 1969) and 2n= 34 is not unknown in the family. It is presumed that this base number (17) has arisen as a result of a cross between progenitors with different base numbers. Bhatt. pollinations won’t occur. Directory of Indian economic plants.7. B. compacted inflorescences and bluish coloured flowers). Twenty nine species belonging to this family are with 2n=34 chromosomes (Federov 1969). Beautiful flowers of Kashmir. G. It is interesting to note that the genus Veronica and Picrorhiza have been placed in the same subtribe (Veroniceae). The capsule size of open pollinated spikes was 7. With 2n=34 chromosomes observed in the present study. 1934. perfect flowers. perianth differentiated into calyx and corolla.3).
O.. 1969. D. 1951. L. In: Supplement to cultivation and utilization of medicinal plants (Eds. and Sharma. Singh. Handbook of ayurvedic medicinal plants. K.. and Tarun Kumar.. International Journal of Pharmacognosy 31:217-222. a source of the Ayurvedic drug Kutaki. Gupta. Rai. Chromosome number of flowering plants. 1996.. pp. A. Studies on medicoethnobotany. G. K. K. Uphof. Florida: CRC Press..K.. O. and Sastry. K. and Maheshwari.K.. 1999 . Red Data Book of Indian Plants. Prajapati. 1959. 350-351. Handa. 2003. Stebbins. 246290. B. M. Danske Biologiske Meddelelser 18: 1-48. A. Jain. Purohit. M. M. D. 2000. Kapoor. C. Chromosome evolution in higher plants.2006.K. Prasad. J. Sharma. domestication and utilization of Picrorhiza kurroa. 1971. Conservation threats to some important medicinal plants of the Sikkim Himalaya. Singh. Sarang. 1885.. Srivastava. 1990. and Banerjee. T. On Pollination in the extremely hot air at Timbuctu. 396-397. Jodhpur: Agrobios. P. diversity. 1996.Quantity of important medicinal herbs collected in Chamoli Garhwal. Jammu: RRL.Y. Picroliv . P. pp. Hooker. 43 .in spite of rain and poverty of insects.. and Kaul. Kapahi. B. A.). Nayer. Kaul. D. G.. West Germany: Ottokaeltz Science Publishers. Dictionary of economic plants. N. New York : Hafner Publishing Co. A. K. B. Madhavan. S. CSIR. Singh. Khajuria. Seth. Description of Picrorhiza kurroa.a natural product protects cells and regulates the gene expression during hypoxia/reoxygenation. Hagerup. 670-684. A handbook of medicinal plants: A complete source book. K. Dansk Bot. L. Gaddipati. P. R. 1993. and Sarin.. Biological Conservation 93: 27-53. London :Edward Arnold. M. 1932. S.. Molecular and Cellular Biochemistry 194: 271-281. Phytotherapy Research 7: 402-407. Sharma.L. K. Hagerup. S. D. Delhi :Bishen Singh Mahendra Pal Singh. S. A. Sindhu. pp. S. P. Flora of British India. E. Ark 8: 120. B.N. L.OAJMAP (2010) Federov. 1987. 333348. Calcutta: Botanical Survey of India. S. pp. G. picroside-1 and kutkoside from Picrorhiza kurrooa . J. Antiinflammatory activity of the iridoids kutkin. R.K. Pollination in the Faeroes . MFP News 6: 12.. pp. S. and Kaul. S. J. T.
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