Survey
* Your assessment is very important for improving the workof artificial intelligence, which forms the content of this project
* Your assessment is very important for improving the workof artificial intelligence, which forms the content of this project
Plant reproduction wikipedia , lookup
Glossary of plant morphology wikipedia , lookup
Gartons Agricultural Plant Breeders wikipedia , lookup
Ecology of Banksia wikipedia , lookup
Ficus macrophylla wikipedia , lookup
Perovskia atriplicifolia wikipedia , lookup
Ornamental bulbous plant wikipedia , lookup
ISSN 2449-8955 European Journal of Biological Research Research Article Seedling establishment in Scaligeria stewartiana (Nasir) Nasir (Apiaceae) from North-West Himalayas, India Kulvinder Kour1, B. L. Bhellum1*, I. A. Hamal2 1 Department of Botany, Govt. College for Woman Parade, Jammu - 180001, Jammu & Kashmir State, India BGSB University, Rajouri, Jammu & Kashmir State, India *Corresponding author: B. L. Bhellum; E-mail: [email protected] 2 Received: 25 April 2016; Revised submission: 10 June 2016; Accepted: 21 June 2016 Copyright: © The Author(s) 2016. European Journal of Biological Research © T.M.Karpiński 2016. This is an open access article licensed under the terms of the Creative Commons Attribution Non-Commercial 4.0 International License, which permits unrestricted, non-commercial use, distribution and reproduction in any medium, provided the work is properly cited. DOI: http://dx.doi.org/10.5281/zenodo.56106 ABSTRACT 1. INTRODUCTION The present paper deals with the morphological diversity of seedling establishment in different populations of Scaligeria stewartiana (Nasir) Nasir, inhabiting in North-West Himalayas. The seedling at different stages in the population of the species revealed marked variability. Seed treatments for germination resulted in understanding of seed dormancy and time required for the seedling establishment. The species develops variability in the genetic level that is transmitted and reflected in the form of morphology in order to adapt itself in the extremes of biotic and abiotic variables. Photographs of the seedling and data compiled in tables catch the specific stages of their seedling development after germination as well variability of the chromosomes. The initiation of tuber formation under the ground level in the soil and allocation of resources perform indispensable role in the life span as well as in paving the way for survival, conservation and distribution of the species. Scaligeria stewartiana (Nasir) Nasir (Tribe pyramidoptereae, Apiaceae) has undergone numerous nomenclatural changes both at genus and species level. Currently, the number of species has been reduced to three. The adaptation of living organisms to the ecological amplitude of any habitat depends upon their genetic composition, which in turn is controlled by various components of their genetic system and finally translated and expressed in the form of morphology. Members of the family Apiaceae have exploited several strategies in order to achieve tremendous diversity in their habitat and geographical distribution in the Himalayas that support rich umbellifer diversity [1]. Scaligeria stewartiana (Nasir) Nasir was recorded from Shiwaliks in Jammu region as a new record for India [2]. The species inhabits isolated populations and turned out to be a diploid, having 2n=20 besides carrying 1-4 extra chromosomes [3]. It is locally known as Jangli Ajwain, the fruit of the species is used as carminative and spice. Plant has thrived high for its conservation by naturally allocating most of its resources towards below ground part, tuber for its survival. Another threat towards its diminishing population is its long period of seedling establishment which takes about four years to establish and a seed dormancy period of full one year. Keywords: Seed germination; Seedlings; Seed set; Tuber; Scaligeria stewartiana; India. European Journal of Biological Research 2016; 6 (3): 145-151 146 | Kour et al. Seedling establishment in Scaligeria stewartiana from North-West Himalayas, India The period of production of seeds year after year is also restricted exactly to four years in which the environmental disturbances, effect of local activities has imposed a lot on its survival. The seeds as well as the leaves are used for various types of gastrointestinal disorders by the locals. The seeds are also used for treatment of digestive disorders of live-stock. 2. MATERIALS AND METHODS Scaligeria stewartiana (Nasir) Nasir of the family Apiaceae is growing in wild situations of different parts of Jammu Province expanding to the different locations at a distance of over 100 Kms, ranging in altitude from 600 to 1200 mts. The sources of the tubers are confined to different localities namely Mansar, Tikri and Reasi. The tubers were transplanted in the botanical garden. Seeds were collected from the different populations and were treated for their germination. Seed germination has been studied under laboratory as well as in the experimental plots of the University Botanic Garden. The seeds collected during May 1995, were treated with 0.1% mercuric chloride for two minutes. Thereafter, the seeds were given a wash in 70 percent alcohol. All traces of mercuric chloride and alcohol were removed by washing the seeds thoroughly with distilled water. For germination in the laboratory, the seeds were divided into replicates of 50 seeds each and then subjected to following physical and chemical treatments. a) Chilling at ± 50C for 15, 20, 30 and 50 days using refrigerator b) Treatment with conc. sulphuric acid for two minutes. c) Hard water treatment for 2 minute. d) Seed coat puncturing e) Seed coat scarification. In all the above treatments including control, no seed germination was recorded under laboratory conditions. However, from the seeds collected during May, 1995, 53 seeds germinated in the experimental plots of the Botanic Garden during January 1997, after sowing during October 1995. There was no germination of the seeds from October 1995 to December 1996. The germinated seeds formed well-established seedlings and some of these seedlings developed into mature plants and flowered in the fourth year. The critical period starts even before germination with predation of seeds, followed by the germination phase that usually allows only less than half of the viable seed to viable seed to emerge, predation on germination and seedlings.[4] Seed germinator was also used in order to germinate seeds. However seeds did not respond to the different treatments of variable temperature provided to facilitate germination. Observation of Botanical Garden population where tubers were transplanted three years earlier, seedlings were observed growing. The measurements of seedlings were made and data was collected. All photos were done by using Olympus PM6 35 Camera. 3. RESULTS 3.1. Observations The seeds from natural population growing at Mansar were collected and treated for germination and showed dormancy of one year (Fig. 1). The germination of seed after dormancy resulted in the formation of radical, which is first to emerge and grow geotropically. After the emergence of the radical two cotyledons are formed which on reaching above ground start turning green. Both the cotyledons were unequal in size in majority of the seedlings and remain green for 20 to 30 days. Seedlings attained a length of 5.0-5.4 cm and the primary root also grow in size which varies from 3.0-3.4 cm. The elongation of primary root resulted in initiation of a tuber in the form of a small swelling at the sub apical position. However, there is no shoot formation. At the end of the growing season the cotyledons wither and the seedlings perennate through the tuber. The epidermis of the tuber starts turning light brown. As the tuber sprouts during next growing period, a basal leaf emerges from tuber above ground (Fig. 1 and Fig. 2 A-B). The basal leaf of the second year seedlings remained green for 30-40 days. During this period photosynthesis in leaves provide nutrition to developing tubers, which attained a size of 8-9.5 mm. The leaves again withered towards the end of the growing season, however, tuber perennated in the soil and it penetrates in the soil up to 15 cm deep European Journal of Biological Research 2016; 6 (3): 145-151 147 | Kour et al. Seedling establishment in Scaligeria stewartiana from North-West Himalayas, India below the ground level. The tubers attain almost the normal size and perennate for the next year (Fig. 2 C). During third year growing season, the tuber sprouted again which resulted in the formation of 2-3 basal leaves as shown in (Fig. 2 D). After further growth the basal leaves emerged above the soil and turned green. The mature leaves formed during third year are larger in size more dissected and remained above ground for 35-45 days. All the resources gathered during growing season were allocated towards below ground tuber [5]. Figure 3. Scaligeria stewartiana (Nasir) Nasir: a matured tuber exposed to show under the ground level. Figure 1. Population of Scaligeria stewartiana (Nasir) Nasir in Mansar. Figure 2. Photographs of seedling of Scaligeria stewartiana (Nasir) Nasir showing different stages A-C: Seedlings at first year after seed germination; D - Second year seedling showing one basal leaf arising from the tuber; E - Third year seedling showing more than one leaf; F - Fourth year tuber showing numerous basal leaves above ground. The tuber that was rounded in the first and second year of growing season attains irregular shape. The leaves after the completion of the growing season wither and tubers enter into dormancy. The tuber which is irregular in shape attaining brownish to black colour in its outer covering again perennates for fourth year and sprouted in the forthcoming season of growth. This fourth year tuber produced 5-9 basal leaves (Fig. 2 E-F and Fig. 3) and their size varied from 10-12 cm. The dissections of the leaves increased and the size of the ultimate leaf segments were observed to be variable. During fourth year of growing season, the tuber turned darker in color and attained diameter up to 3.5 cm. The size of the fourth year tuber was comparable to the size of the tuber transplanted from natural populations. During the same year (4th years) these tubers initiated shoot development, which subsequently developed flowers in the orders starting from primary to higher orders. 3.2. Seed morphology Like all umbellifers, the species studied now, is characteristic in having a schizocarpic fruit; the seed coat is fused with the fruit wall. The fruit is the dispersal unit, which on maturity splits into two European Journal of Biological Research 2016; 6 (3): 145-151 148 | Kour et al. Seedling establishment in Scaligeria stewartiana from North-West Himalayas, India mericarps. Each mericarp alongwith the seed forms a single dispersal unit. The fruit is subdidymous, broad at base and narrow towards apex where it is capped by a stlypodium from which two recurved styles emerge. Each mericarp carries five primary ridges, one dorsal, two commissural and two intermediate. The ridges are corky, obtuse and run parallel to the longitudinal axis of the mericarp. In between the primary ridges, are present vallecular spaces. Seed weight: The seeds are small in size; each mericarp measures 1.5-2.3 (x̄ 2.1) mm in length and 0.8-1.1 (x̄ 0.09) in breadth, ( commissural face). The seed weight of 100 mericarps varies between 70-95 (x̄ 90) mg. percent in Tikri and x̄ 60.2 ± 0.51 (0-65.0) in Reasi population. The fruit set was also recorded separately for normal disomics and those bearing 1-4 extra chromosomes, beside the norma complement (Table 1). The data given in Table 18 reveals that in comparison to normal disomics, the percentage fruit set reduces, with increase in the number of extra chromosomes, in all the umbel orders. In primary umbel it reduced from 93.47 in disomics to 52.13 percent in plants with 2n=20 ± 4 extra chromosomes. Similar trend was observed in the fruit set of secondary, tertiary and quarternary umbel orders. 3.3. Seed set As the seed germinate, the radical is the first to emerge and grow geotropically. Subsequently, the two cotyledons emerge and on reaching above ground (Fig. 2 A-B) start turning green. The cotyledons are unequal sized in majority of the seedlings and remain green for 20-30 days. During this period the seedlings attain a length of 5.0-5.4 cm. The primary root also grows in size, which varies from 3.0-3.4 cm. As the elongation of primary root is taking place; the development of a tuber is initiated in the form of a small swelling at its sub-apical position (Fig. 2 C). However, there is no shoot formation. Towards the end of the growing season, the cotyledons wither and the seedlings perennate through the tuber. The epidermis of the tuber turns light brown. As the tuber sprouts during next growing period, a basal leaf emerges from the tuber and reaches above ground (Fig. 2 D). Although all seedlings at the stage carried a single basal leaf but the dissection of the leaf was variable. The basal leaf of these second seedlings remained green for 30-40 days. During this period, photosynthesis in leaves provided nutrition to developing tubers, which attained a size of 8 to 9.5 mm. The leaf withered towards end of the growing period and the tuber perennated in the soil, upto 15 cm deep from the ground level. During third year growing season, the tuber again sprouted and formed 2-3 basal leaves (Fig. 2 E-F). After further growth, the basal leaf emerges above the soil and turned green. The mature leaves formed during third year were more dissected and large in size (9-10 mm). Scaligeria stewartiana (Nasir) Nasir exhibits variation in seed set at intra- as well as at inter population level. The umbels of different order portray variation in their contribution to the total fruit yield of an individual plant. In Mansar population, the tertiary umbels contribute maximum to the total fruit yield of the individual plant. On the contrary secondary umbels are the major contributors to the fruit yield per plant in Tikri and Reasi populations. However, irrespective of the contribution of different umbel orders to the total fruit yield, the fruit set decreases from primary to the higher order umbels in all the three population. The average fruit set in primary umbel of all the three populations was above 90 percent. It was x̄ 90.62 ± 0.55 (77.7-100) percent in Mansar population, x̄ 93.95 ± 0.48 (75.0-100) percent in Tikri population and x̄ 93.38 ± 0.26 (80.0-100) percent population. Percentage fruit set in secondary order umbels varied between 88-90 percent, being x̄ 88.9 ± 0.35 (72.0-100) in Mansar, x̄ 88.48 ± 0.32 (81.895.0) in Tikri, x̄ 90.94 ± 0.23(79.0-100) percent in Reasi population. The fruit set in third order umbels ranged between 70-83.38 percent. The Mansar population had 70.31 ± 1.49 (49.6-100) percent whereas in Tikri and Reasi populations, fruit set was x̄ 83.38 ± 0.71 (70-100) and x̄ 81.24 ± 0.36 (36.2-90) percent respectively. There was wide variation in the fruit set of fourth order umbels. It was x̄ 65.72 ± 1.36 (0-90.6) percent in Mansar, x̄ 78.1 ± 0.49 (0-88.8) 3.4. Seedling establishment European Journal of Biological Research 2016; 6 (3): 145-151 149 | Kour et al. Seedling establishment in Scaligeria stewartiana from North-West Himalayas, India Table 1. Fruit set at umbel in Scaligeria stewartiana (Nasir) Nasir. Primary order Chromosome number (2n) 20 20+1 20+2 20+3 20+4 Number Percent fruit set Bisexual Fruit Flower formed x̄ 160± 149± 93.47± 1.31* 1.21 0.24 (110(91(82.6179)** 179) 100) 159± 129± 81.43± 6.6 5.35 1.91 (136(104(76.5163) 144) 88.8) 138± 87± 63.25± 12.4 7.81 5.90 (126(90(72-81) 225) 182) 132± 80± 60.1± 9.6 6.5 4.91 (125(65(52-70) 169) 118) 130± 67± 52.13± 8.9 4.2 3.24 (120(54(45-80) 175) 140) Secondary order Number Bisexual Fruit Flower formed 340± 326± 9.25 4.27 (170(152660) 660) 370± 287± 8.93 6.91 (175(121710) 642) 410± 195± 10.68 5.08 (190(81690) 340) 390± 180± 11.62 5.36 (180(72560) 276) 480± 227± 11.58 5.47 (225(86720) 175) Tertiary order Percent fruit set 95.94± 0.28 (89.9100) 77.6± 0.83 (7090.47) 47.8± 0.54 (42.850.0) 46.2± 0.53 (40.149.2) 47.3± 0.63 (38.249.4) Number Quarternary order Number Percent Bisexual Fruit fruit set Bisexual Fruit Flower formed Flower formed 396± 180± 72.23± 179± 127± 5.95 3.21 2.34 3.92 2.39 (250(120(48(0(0-275) 450) 490) 100) 253) 352± 126± 35.8± 171± 60± 7.43 2.46 2.0 4.23 2.47 (240(53(22.2(0(0-269) 560) 350) 62.5) 183) 360± 274± 45.15± 79± 29± 6.38 2.87 2.50 2.31 0.85 (255(117(28.6(0(0-182) 610) 360) 50) 124) 315± 77± 24.63± 69± 24± 5.27 4.21 9.6 2.11 0.74 (240(42.0(17.6(0-169) (0-61) 560) 280) 50) 290± 65± 22.59± 82± 28± 4.23 0.95 4.24 2.6 0.9 (210(32(15.2(0-159) (0-84) 510) 229) 45) Percent fruit set 71.2± 1.32 (4092.3) 35.0± 2.0 (067.9) 37.2± 2.3 (068.1) 35.2± 3.1 (069.5) 34.0± 3.43 (052.9) Table 3. Dry biomass of Scaligeria stewartiana (Nasir) Nasir above and below ground parts of the seedings at different stages of development. Total dry Dry biomass of below Dry biomass of above Seedings (N) biomass (mg) ground parts (mg) ground parts (mg) Ratio b:c stage a b c x̄ 0.2330±0.0005* 0.1156 ± 0.0003 0.1174±0.0005 Ist year 20 1.01 (0.2021-0.2497)** (0.1039-0.1251) (0.1094-0.1279) 0.2286±0.0006 0.1137±0.0005 0.1149±0.0007 IInd year 20 1.01 (0.1935-0.2481) (0.1120-0.1296) (0.1012-0.1256) 0.1353±0.0003 0.1190±0.0005 0.1163±0.0006 IIIrd year 20 0.97 (0.1151-0.1532) (0.1036-0.1295) (0.1020-0.1347) 0.2570±0.0005 0.1230±0.0006 0.1110±0.0005 IV year 10 0.90 (0.2193-0.2695) (0.936-0.1435) (0.996-0.1239) * Mean, S.E ** Range These remained green and above ground for 35-40 days. During this period, the tuber also grew in size upto 1.5 cm in diameter. During this period, tuber also developed adventitious roots. The tuber, which was almost globular during first and second year, assumed irregular shape in the third year. Towards the end of the growing season, the leaves withered and tubers again entered into dormancy. After perennation of the third year, the tubers sprouted again during the growing season of fourth year. The new basal leaves emerged above ground. Each tuber produced 5-9 basal leaves (Fig. 2 A-B), which varied 10.2-12.5 cm in length. The dissections of the leaves increased and the size of ultimate leaf segments were also variable. The tuber became darker in colour and attained diameter upto 3.5 cm. The number of adventitious roots emerging from each tuber also increased. The size of the fourth year tuber was comparable to the size of tubers transplanted from natural populations. During the European Journal of Biological Research 2016; 6 (3): 145-151 150 | Kour et al. Seedling establishment in Scaligeria stewartiana from North-West Himalayas, India same year (4th year) these tuber initiated shoot development, which subsequently produced flowers in the same order as was observed in case of tubers transplanted from natural populations. 3.5. Seed viability The viability of the seeds was determined by tetrazolium test (Tz-test). The seeds of various populations exhibited variation in their percentage viability. In all the populations, seed viability was above 85 percent (Table 4). The strategy of resource allocation in the species has been assessed after analyzing dry biomass at various stages of seedling establishment and seed set. The different stages of seedling exhibit distinct morphology. In the subsequent years, seedlings produce shoots and umbels of different orders. The dry biomass of the seedlings at different stages of development is represented in the Table 3. Table 4. Seed viability in Scaligeria stewartiana (Nasir) Nasir. Seeds Seeds Population Percentage tested viable Botanic 95 89 93 Garden Mansard 100 91 91 Tikri 120 102 85 Reasi 90 85 94.4 4. DISCUSSION The life history pattern of a species is the result of evolutionary decision in order to achieve successful establishment, adaptation and future evolutionary diversification [6]. The perpetuation of a species depends at the rate at which individuals of population contribute to future generations, which in turn is influenced by the number of reproductive attempts during its life time, age of first reproduction and reproductive rate. The herbaceous perennial Scaligeria stewartiana (Nasir) Nasir produces leaves and flowers every year from the underground tuberous root which however it is not an organ of vegetative multiplication [7]. The observations on seed biology of the species have revealed that the seed undergoes germination after attaining dormancy for the period of one year. After germination of seeds the seedlings get established in four years, after that every individual plant flowers for 4 to 5 years before perishing. The reproductive output also increases with age, as the tuber increase in size. As compared to only two umbel orders during first year of reproductive age, the plant in the subsequent years produce umbels upto fourth order [8]. During first two years of seedling establishment, the species invests more resources (dry biomass) in the formation of aerial parts and in third and fourth year, the investment is more in the production of tuber. Long gestation period in seedling establishment enables the species to reduce competition and maintain genetic composition of the populations. The weather conditions are main cause of mortality. However, permanent snow cover can protect seedling from frost and heaving injuries [9-11]. 5. CONCLUSION Figure 4. Scaligeria stewartiana (Nasir) Nasir: representing comparison of seed viability at different locations in Jammu and Kashmir. The populations of the species are decreasing due to its seed dormancy and other types of environmental disturbances, natural and human interferences. The seed to seed cycle of the species takes four years for completion whereas the total life span of the species ranges from 7-8 years. During first two years of seedling establishment, the species invest more resources (dry biomass) in the formation of ariel parts and in 3rd and 4th year, the investment is more in the production of tuber. The gestation period reduces competition between European Journal of Biological Research 2016; 6 (3): 145-151 151 | Kour et al. Seedling establishment in Scaligeria stewartiana from North-West Himalayas, India established individual and fresh recruits. However, the species needs special attention due to a number of threats. 3. Kour K, Hamal IA, Gupta SK. Chromosomal variability in Scaligeria stewartiana (Nasir) E. Nasir. J Cytol Genet. 1992; 27: 43-45. 4. Willson MF. Plant reproductive ecology. John Wiley and Sons, Inc., New York, 1983. 5. Koul V. Resource allocation in relation to floral structure and breeding system in some member of commelinaceace. Ph.D. Thesis, University of Jammu, 1998. 6. Putz N, Sakkau I. Seedling establishment bud movement and subterranean diversity of subterranean system in Apiaceae. Flora. 2002; 197: 385393. 7. Constance L, Chuang T, Bell CR. Chromosome numbers in Umbelliferae. Am J Bot. 1971; 58(6): 577-587. 8. Mukerjee PK, Constance L. Umbliferae of India. Oxford and IBH. Publishing, New Delhi, 1993. 9. Bergsten U, Nilsson JE. Early seedling growth of Pinus sylvestris (L.) after sowing with a mixure of stand and orchard seed in dence spacings. Can J Forest Res. 2001; 31(7): 1184-1194. ACKOWLEDGEMENTS Authors are grateful to Prof. A. K. Koul, Dean Academics, BGSB University, Rajouri for helpful suggestions and thankful to Principal Govt. College for Women, Parade Jammu for laboratory facilities. AUTHORS’ CONTRIBUTION KK: Data collection, Interpretation, Writing, Discussion; BLB: Manuscript designing, Discussion; IAH: Guidance, Interpretation. The final manuscript has been read and approved by all authors. TRANSPARENCY DECLARATION The authors declare no conflicts of interest. REFERENCES 1. Sarukhan J. Demography problems in tropical system. Bot Monogr. 1980; 15: 161-188. 2. Hamal, I. A. Cytotaxonomy of the umbellifers of Kashmir Himalayas. Ph. D. Thesis. Jammu University, 1981. 10. Bergsten U, Nilsson JE. Effects of seed weight and seed type on early seedling growth of Pinus sylvestris under harsh and optimal conditions. Scand J Forest Res. 2002; 17: 118-130. 11. Vernet P, Harper JL. The costs of sex in seaweeds. Biol J Linn Soc. 1980; 13: 129-138. European Journal of Biological Research 2016; 6 (3): 145-151