Seed germination behavior of Tsuga dumosa (D. Don) Eichler (Himalayan Hemlock), a rare gymnosperm of Indian Himalayan Region.

Seed germination behavior of Tsuga dumosa (D. Don) Eichler (Himalayan Hemlock), a rare gymnosperm of Indian Himalayan Region. | Authorea try { document.documentElement.classList.add('js'); } catch (e) { } var _gaq = _gaq || []; _gaq.push(['_setAccount', 'G-8VDV14Y67G']); _gaq.push(['_trackPageview']); (function() { var ga = document.createElement('script'); ga.type = 'text/javascript'; ga.async = true; ga.src = ('https:' == document.location.protocol ? 'https://ssl' : 'http://www') + '.google-analytics.com/ga.js'; var s = document.getElementsByTagName('script')[0]; s.parentNode.insertBefore(ga, s); })(); Skip to main content Preprints Collections Wiley Open Research IET Open Research Ecological Society of Japan All Collections About About Authorea FAQs Contact Us Quick Search anywhere Search for preprint articles, keywords, etc. Search Search ADVANCED SEARCH SCROLL This is a preprint and has not been peer reviewed. Data may be preliminary. 18 July 2025 V1 Latest version Share on Seed germination behavior of Tsuga dumosa (D. Don) Eichler (Himalayan Hemlock), a rare gymnosperm of Indian Himalayan Region. Authors : Abhinay Bhardwaj 0009-0004-8594-3530 [email protected] , Manisha Thapliyal [email protected] , and Sheeshram Dangwal Authors Info & Affiliations https://doi.org/10.22541/au.175283114.49903034/v1 360 views 132 downloads Contents Abstract Supplementary Material Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract Large varieties of conifer species have found their origin in the Himalayas, possessing high ecological importance, silvicultural significance, and economic value. Tsuga dumosa is one such species of the Indian Himalayan Region that is less known among researchers. It is classified as “Least Concern” in terms of IUCN status. So, it becomes very important to manage the species scientifically in an organized and sustainable manner. The present study contributes to the study of various aspects of seed biology, seed characteristics, and germination behavior of Tsuga dumosa to support its propagation and scientific management. The study was conducted at the Forest Tree Seed Laboratory, Silviculture Division, FRI, Dehradun, from February to June 2022. The cones of Tsuga dumosa were collected from Tilthin, Narayanashram, Dharchula, Pithoragarh, and Uttarakhand, and seeds were extracted in the Seed Processing Unit, FRI Dehradun. The seed germination experiment was laid out in a Completely Randomized Design (CRD) with 3 replicates. Data were recorded on seed weight, seed length, seedling length, moisture content, and germination response to various pretreatments. Additionally, the use of the right growth promoter in the appropriate concentration on germination enhancement was evaluated. Among the treatments, seeds soaked in 300 ppm GA3 for 24 hours followed by stratification for 30 days showed the highest germination percentage. For seedling growth, seeds stratified for 45 days and sown in vermiculite medium exhibited superior performance, as better root length, shoot length, and collar diameter were recorded. Seed germination behavior of Tsuga dumosa (D. Don) Eichler (Himalayan Hemlock), a rare gymnosperm of Indian Himalayan Region. Abhinay Bhardwaj 1* , Manisha Thapliyal 1* , Sheeshram Dangwal 2 Forest Tree Seed Laboratory, Silviculture and Forest Management Division, Forest Research Institute, Dehradun, 248006, Uttarakhand, India ABSTRACT Large varieties of conifer species have found their origin in the Himalayas, possessing high ecological importance, silvicultural significance and economic value. Tsuga dumosa is one such species of the Indian Himalayan Region that is less known among researchers. It is classified as “Least Concern” in terms of IUCN status. So, it becomes very important to manage the species scientifically in an organized and sustainable manner. The present study contributes to the study of various aspects of seed biology, seed characteristics and germination behavior of Tsuga dumosa to support its propagation and scientific management. The study was conducted at the Forest Tree Seed Laboratory, Silviculture Division FRI, Dehradun, from February to June 2022. The cones of Tsuga dumosa were collected from Tilthin, Narayanashram, Dharchula, Pithoragarh Uttarakhand and seeds were extracted in the Seed Processing Unit, FRI Dehradun. The seed germination experiment was laid out in a Completely Randomized Design (CRD) with 3 replicates. Data were recorded on seed weight, seed length, seedling length, moisture content and germination response to various pre-treatments. Additionally, the use of right growth promoter in the appropriate concentration on germination enhancement was evaluated. Among the treatments seeds soaked in 300 ppm GA 3 for 24 hours followed by stratification for 30 days showed the highest germination percentage. For seedling growth, seeds stratified for 45 days and sown in vermiculite medium exhibited superior performance, as better root length, shoot length and collar diameter were recorded. Key words: Tsuga dumosa , seed, dormancy, cold stratification, Himalayan conifers, germination, GA 3 treatment, forest regeneration Authors: Abhinay Bhardwaj* ( [email protected] ), Manisha Thapliyal* ( [email protected] ), Sheeshram Dangwal- Forest Tree Seed Laboratory, Silviculture and Forest Management Division, Forest Research Institute, Dehradun-248006, Uttarakhand, India INTRODUCTION Tsuga dumosa , commonly known as Himalayan Hemlock or locally as “Tansin”, is a conifer species native to the Indian Himalayan Region, particularly in the Kali Valley of Pithoragarh district, Uttarakhand. It also occurs in parts of Nepal, Bhutan, and southeastern Tibet. In India, its presence and natural regeneration have been documented in the Munsiyari and Dharchula ranges of Pithoragarh Forest Division (ICFRE, 2013), as well as in the East Almora Division (Osmaston, 1927). The species grows predominantly along the southern slopes of the Eastern Himalayan range, stretching from Kumaon to the inner valleys, at elevations between 2,100 and 3,000 meters above sea level (Miehe et al., 2015). According to forest records, it covers approximately 4,607 ha of reserve forest and 230 ha of protected forest in the Pithoragarh division, with an estimated population of 11,500 trees (Chandran, 2011). Based on Champion and Seth’s (1968) classification, T. dumosa falls under the East Himalayan Dry Temperate Coniferous Forest type. It is usually found in association with Pinus wallichiana , Abies densa , and Rhododendron spp., but may also form pure stands in certain locations. These forests are categorized as cool, humid coniferous ecosystems with a mixed understory of evergreen and deciduous broadleaf species (Wangda & Ohsawa, 2006). Quantitative ecological studies in the region have identified T. dumosa as a dominant species, with the highest recorded Importance Value Index (IVI) of 80.07 among coexisting flora (Naithani, 2018). Despite its ecological prominence, T. dumosa exhibits regeneration challenges. Dhungana & Chetri (2024) reported a ‘fair’ seedling status at mid to lower elevations in central Nepal, which declined to ‘poor’ in upper ranges due to factors such as livestock grazing and recurring forest fires. However, in certain compartments like Shoble No. 7 of Dharchula Range, trees of T. dumosa have been recorded with heights up to 44 m and girths of 4 m, indicating favourable site conditions for growth and regeneration (Chandran, 2011). From a silvicultural perspective, hemlocks are slow-growing, long-lived, and highly shade-tolerant compared to other conifers, but they are also drought-sensitive (Havill et al., 2008). In comparative studies on shade tolerance, T. dumosa was classified as the most shade-tolerant species among Himalayan conifers (Gratzer et al., 2004). Taxonomically, the genus Tsuga (family Pinaceae) includes 8–10 species—four from North America and four to six from eastern Asia. Earlier botanists grouped hemlocks with firs, pines, or spruces due to morphological similarities, but Carrière later placed them under a distinct genus Tsuga (Troup, 1921). Reproductive structures in conifers such as T. dumosa are cone-based, with seeds derived from naked ovules (Willan, 1985). Most conifer seeds possess adnate or articulate wings, aiding in wind dispersal (Khullar et al., 1991). Hemlocks, like Abies and Cedrus , also have resin vesicles in their seed coats (Singh, 1978). Cones disintegrate upon reaching maturity, a characteristic that complicates seed collection timing. To ensure viable seed harvest, cones are often collected just before full maturity and subjected to post-harvest ripening (Thapliyal, 2014). Seed dormancy in conifers is a key challenge affecting natural regeneration. Successful regeneration depends on factors such as seed production, dispersal, germination, and establishment (Thapliyal, 2014). Hemlocks disperse seeds slowly—over 3 to 12 months—as cones gradually release seeds. Laboratory studies on Tsuga heterophylla have reported shallow dormancy and intermediate storage behaviour, with an optimal seed moisture content of 9–12% for storage (Gosling, 2007). In addition to ecological importance, T. dumosa possesses medicinal potential. Essential oils extracted from its tissues exhibit antibacterial and antioxidant activities, attributed to bioactive compounds such as tannins, alkaloids, terpenoids, and flavonoids (Ojha & Joshi, 2023). The present endeavour is toward assessing the various processes and morphological parameters involved in determining the seed characteristics and germination behaviour of Tsuga dumosa . Duration of moist chilling with growth promoters will be worked out for overcoming dormancy. Understanding the dormancy-breaking requirements and early seedling growth under controlled conditions can help optimize nursery protocols and enhance field planting success for this lesser-studied conifer of the Indian Himalayan Region. MATERIALS AND METHODS Seed Collection and processing Mature cones of Tsuga dumosa were collected from within its natural distribution range in Tilthin, Pithoragarh Uttarakhand (Figure 1). 20 mature and healthy trees were selected randomly for cone collection at least 150 m apart from each other to reduce genetic similarity. For optimal laboratory conditions Forest Tree Seed Laboratory of FRI, Dehradun was chosen. Geographic information of the collection site and experiment site is provided in (Table 1). The collected cones were sun-dried under ambient conditions at the premises of Seed Processing Unit, Forest Research Institute for nearly 2 weeks (Figure 2). Seeds were then extracted from the cones (Figure 3), cleaned, dewinged, and stored in sealed containers at 10°C and 15% relative humidity. Seed storage protocol followed low-temperature, low-humidity conditions to ensure seed viability. Prior studies have shown that conifers seeds such as Pinus roxburghii , exhibit enhanced storability when kept in sealed glass jars at 5±1°C (Gautam et al., 2005). Cone specific gravity is also considered an important seed maturity indicator in conifers like Pinus kesiya , where lower gravity has been correlated with higher germination (Singh & Kachri, 2006). Seed characteristics Seed Weight: Seed weight is generally expressed as the weight of 1000 pure seeds but due to availability of smaller number of seeds weight of 500 pure seeds was taken. Total weight by counting of 500 seeds was taken and then weight of 5 Replicates of 100 seeds each were taken on the basis of which mean and standard deviation was calculated. Number of seeds in 1g was also calculated (Table 2). Seed Length and Width: Length and breadth of the seeds (with wing and without wing) was observed using a ruler/scale because seeds were too small to be recorded in an electronic caliper. 4 replications of 25 undamaged seeds (with wing and without wing) were taken and measured for their length and breadth on the basis of which mean and standard deviation was calculated (Table 3). Moisture Content Determination: Moisture content was determined using aluminum moisture container box enclosed with aluminum foil. Approximately 0.20 g of seeds per replicate (four replicates) were dried in aluminum moisture containers at 103°C ± 1 for 17 hours. Further drying and desiccation of seeds was done to determine the moisture content. Germination experiment: Initially germination test was done without any pretreatment to evaluate baseline performance. To break the dormancy the collected seeds of Tsuga dumosa were subjected to cold stratification at 5°C in a moist towel paper. Temperatures below freezing was avoided as it may injure the imbibed seeds. Germination testing using 300ppm concentration of GA 3 was also done. (Figure 4). Laboratory germination tests were carried out in 10 cm diameter. Petri plates lined with moistened blotters maintained at 20 ± 3°C. Whenever moisture loss was detected, distilled water was sprinkled. There were four treatments in this experiment including the control. Three were conducted in Petri plates and one using expanded vermiculite (mica) medium. Experimental Design and Evaluation: The experiment was laid out in CRD (Completely Randomized Design) with 3 replications per treatment and 25 seeds per replicate. A seed was regarded as germinated when the radicle had grown to approximately 1cm in length. Abnormal seedlings were excluded from final counts as they rarely survive to produce a plant. Germination performance was quantified as per the international Seed Testing Association (ISTA, 2010) standards. The parameters evaluated included Germination Percentage (GP), Peak Value (PV), Germination Value (GV), Mean Daily Germination, Vigor Index (VI) and Germination Index (GI) (Table 4). Seedling morphological traits, including root length, shoot length and collar diameter were recorded 28 days after initial germination. Seed Morphometrics The average weight of 500 Tsuga dumosa seeds was 1.49g, corresponding to an average of 331.15 ± 0.97 seeds per gram and 3314.5 ± 5.54 seeds per 10 grams. The mean weight of 10 seeds was recorded at 0.03 ± 0.005g and that of 100 seeds at 0.30 ± 0.01g (Table 2). Seed length and width, with and without wings were measured for 200 seeds. The average length of seeds with wings was 8.8 ± 1.90 mm, while the width was 3.9 ± 0.70 mm. The average length and width of seeds without wings were 3.90 ± 0.70 mm and 2.6 ± 0.70 mm, respectively (Table 3). Germination Response and Seedling Growth Significant variation was observed in seed germination and seedling traits among treatments (Table 4). The control (T 0 ) exhibited the lowest germination percentage (6.67%) with a mean germination time (MGT) of 7.80 days and germination index (GI) of 0.66. Treatment (T 2 ) comprising seeds soaked in 300 ppm GA 3 for 24 hours and stratified for 30 days, resulted in the highest germination percentage (34.66), GI (2.85) and (VI) vigor index (141.79). T 3 involving 45-day stratification and vermiculite medium, recorded germination percentage 25.33%. Seedling parameters (Table 5) showed that T 3 resulted in the highest average root length (1.43 ± 0.16 cm) and collar diameter (0.79 ± 0.03mm), while T 2 exhibited the highest average shoot length (2.77 ± 0.17cm). Statistical Analysis ANOVA (Table 6) indicated that treatment effect on germination percentage was highly significant (F=18.386; p =0.001). The coefficient of variation (CV) was 25.17%. Critical difference (CD) at 5% and 1% levels were 9.477 and 13.788, respectively. Treatments T 2 and T 3 were statistically at par and significantly superior to T 0 and T 1 . * [email protected] 1 Department of Infectious Disease, Imperial College London, Du Cane Rd, London W12 0NN UK. 2 Department of Biotechnology, Hezekiah University Umudi, Imo State, Nigeria. DISCUSSION The present study offers a detailed examination of seed and seedling traits of Tsuga dumosa , with an emphasis on improving germination and early growth through stratification, growth regulators and suitable media. The results reveal that seed parameters such as weight, length and moisture content fall within the range observed in other Himalayan conifers. The average moisture content recorded in this study (16.25%) closely aligns with earlier findings in Abies pindrow where mature seeds ranged from 15.82% to 16.60% (Singh, 1998), indicating physiological maturity at the time of collection. Seed germination response was significantly influenced by pre sowing treatments. Among all treatments, T 2 (30 days stratification+ 300 ppm GA 3 ) produced the highest germination percentage (34.67%), germination index (2.85) and vigor index (141.79). These findings confirm the positive role of gibberellic acid in breaking physiological dormancy, consistent with earlier studies on Picea smithiana (Chandra & Chauhan, 1977) and Abies pindrow (Rawat el al.,2008). Similarly in Tsuga canadensis , unstratified seeds failed to germinate at high temperatures and the highest germination (68%) was achieved at 53°F following 10 weeks of stratification (Olson el al., 1958), indicating that cool stratification is critical for dormancy release in Tsuga species. Although T 2 performed best for germination, T 3 (45- day stratification with vermiculite medium) promoted superior seedling growth, particularly in terms of root length (1.43 cm), collar diameter (0.79mm) and total seedling length (4.13cm). This suggest that while hormonal and chilling treatment facilitate dormancy release and synchronized germination, vermiculite provides a favorable microenvironment for robust root and shoot development. Similar trend was observed in Cedrus deodara, where stratification up to 75 days significantly improved both germination (82.1%) and germination value (19.5) (Sofi & Bhardwaj, 2008). Despite promising results under controlled conditions, germination in nursery settings was poor, likely due to elevated temperatures and site-specific climatic limitations. This is consistent with observations by Troup (1921), who reported delayed and inconsistent germination in Tsuga under natural nursery conditions, often exceeding from 6 weeks to 6 months. These findings emphasize the importance of using controlled environments for the propagation of dormant high-altitude species. Seed size and weight are critical indicators of germination and seedling performance. The results of this study align with prior research demonstrating a positive correlation between seed size and seedling vigor in conifers (Spurr, 1944; Fehr & Weber, 1968). Moreover, variation in germination due to seed source have been widely reported across species such as Pinus brutia (Isik, 1986) , Pinus greggii (Dvorak et al., 1996), Acacia mangium (Salazar, 1989), and Pinus roxburghii (Roy et al., 2004). Environmental factors like temperature, rainfall and soil moisture can also impact seed development and quality (Mughal & Thapliyal, 2013). Genetic variability and environmental heterogeneity within provenances may further explain the variation in seed performance. Overall, the study demonstrates that integrated pre-sowing strategies, particularly the combined use of GA 3 and cold stratification, can enhance germination in T. dumosa , while vermiculite supports healthy seedling development. However, for successful field establishment further research is needed on nursery microclimate management and selection of optimal seed sources. CONCLUSION The present study provides valuable insights into the seed germination behavior of Tsuga dumosa , an ecologically and silvicultural significant conifer of Indian Himalayan Region. These results have direct implications for the conservation and sustainable management of T. dumosa, particularly in the face of habitat degradation and climate change. From the present study it can be concluded that stratification is the most important factor affecting the germination response in conifers. Stratified seeds germinated much better than unstratified seeds and it is very important to break the dormancy of the seeds of conifer species. Right growth promoter in appropriate concentration plays a vital role in enhancing the germination response. According to the study seeds stratified for 30 days after soaking in 300 ppm GA 3 for 24 hours turned out to be the best treatment given to enhance the germination response of seeds of T. dumosa . For growth of seedling T3 was best as it resulted in better root length, shoot length and collar diameter. However, Indian Himalayan Region, particularly dominated by T. dumosa remain among the least studied ecosystems. Limited research has focused on the community composition, regeneration dynamics and seed characteristics of this species. Broader concentration of GA 3 can be used further by the researchers in its germination studies. Study on nursery techniques can be carried out in different agro-climatic zones of Uttarakhand. Seed quality testing can be done more precisely as seed emptiness was an issue in this study. Right seed storage protocols to enhance longevity can prove to be some important factors in utilizing the species to its full potential. Further studies on seedling ecology and field performance will compliment these findings and aid in the effective management, long-term conservation of this species within these unique forest habitats. This will help prevent this jewel conifer species from the threat of getting into critical categories in future. Acknowledgement The authors sincerely acknowledge the Silviculture and Forest management Division, Forest Tree Seed Laboratory, FRI Dehradun for providing optimum laboratory facilities and research support. This paper is a part of Abhinay Bhardwaj’s Master’s Thesis. We would like to express our special thanks to Ms. Shweta Rawat, Dr. Karuna Phular, Mr. Deepak Ghuniyal, Ms. Vaishakhi Prem Chand, Mr. Prabal Kumar who truly helped us in collecting the information regarding the species. * [email protected] 1 Department of Infectious Disease, Imperial College London, Du Cane Rd, London W12 0NN UK. 2 Department of Biotechnology, Hezekiah University Umudi, Imo State, Nigeria. REFERENCES Bhat, H. A., Mughal, A. H., Dar, M. U. D., & Mugloo, J. A. (2018). Cone, seed and germination characteristics in silver fir ( Abies pindrow Spach) along the altitudinal gradient in Western Himalayas. International Journal of Chemical Studies , 6 (2), 2052–2055. Chandra, J. P., & Chauhan, P. S. (1977). 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U.S. Department of Agriculture, Forest Service. (1974). Seeds of woody plants in the United States (Agricultural Handbook No. 450). Vakshasya, R. K., Rajora, O. P., & Rawat, M. S. (1992). Seed and seedling traits of Dalbergia sissoo Roxb.: Seed source variation studies among ten sources in India. Forest Ecology and Management , 48 , 265–275. Wangda, P., & Ohsawa, M. (2006). Structure and regeneration dynamics of dominant tree species along an altitudinal gradient in a dry valley slope of the Bhutan Himalaya. Forest Ecology and Management , 230 , 136–150. Willan, R. L. (1985). A guide to forest seed handling . Food and Agriculture Organization of the United Nations. TABLES Site of collection Uttarakhand Tilthin, Narayanashram, Dharchula, Pithoragarh Forest Division 2678m (8786ft.) 29.86°N 80.52°E Experiment site Uttarakhand Forest Tree Seed Laboratory, Forest Research Institute Dehradun 640m (2100ft.) 30.34°N 77.99°E Table 1. Geographic information of site of collection and experiment site Average no. of seeds in 1gm 331.5±0.97 Average no. of seeds in 10gm 3314.50±5.54 Average weight of 10 seeds (gm) 0.03±0.005 Average weight of 100 seeds (gm) 0.30±0.01 Table 2. Seed weight * [email protected] 1 Department of Infectious Disease, Imperial College London, Du Cane Rd, London W12 0NN UK. 2 Department of Biotechnology, Hezekiah University Umudi, Imo State, Nigeria. Length of seeds (with wing) (mm) 8.8±1.90 Width of seeds (With wing) (mm) 3.9±0.70 Length of seeds (without wing) (mm) 3.9±0.70 Width of seeds (Without wing) (mm) 2.6±0.70 Table 3. Average seed length and width * [email protected] 1 Department of Infectious Disease, Imperial College London, Du Cane Rd, London W12 0NN UK. 2 Department of Biotechnology, Hezekiah University Umudi, Imo State, Nigeria. T0 6.67 7.80 0.67 0.31 0.66 21.20 T1 13.33 6.20 1.71 1.88 1.74 53.86 T2 34.66 11.73 1.75 3.21 2.85 141.79 T3 25.33 10.94 1.51 2.10 2.12 104.62 Table 4. Germination Parameters- T0- Control, T1- 15days stratification, T2- 30 days stratification+0.03% GA 3 , T3- 45 days stratification +vermiculite media T0 1.02±0.21 2.16±0.15 0.67±0.11 T1 1.32±0.19 2.72±0.10 0.72±0.05 T2 1.32±0.17 2.77±0.17 0.77±0.03 T3 1.43±0.16 2.70±0.24 0.79±0.03 Table 5. Seedlings parameters of T. dumosa Source of variation Degrees of freedom Sum of squares Mean sum of squares F cal F prob Treatments 3 1397.333 465.778 18.386 0.001 Error 8 202.667 25.333 - - Total 11 - - - - Coefficient of variation=25.166. Treatments found Significant at 1% and 5% level of significance CD(0.01) = 13.788 CD(0.05) = 9.477 Treatment Average 34.667 25.333 13.333 6.667 Critical Difference (CD) Compared a a b b Table 6. Comparison of Treatment Means with Critical Difference (0.05) GA3 Gibberellic Acid CRD Completely Randomized Design PV Peak Value GV Germination Value GP Germination Percentage MGT Mean Germination Time VI Vigor Index GI Germination Index ANOVA Analysis Of Variance IUCN International Union for Conservation of Nature FRI Forest Research Institute ICFRE Indian Council of Forestry Research and Education USDA United States Department of Agriculture Supplementary Material File (figures of research paper abhinay bhardwaj tsuga dumosa.docx) Download 1.11 MB Information & Authors Information Version history V1 Version 1 18 July 2025 Copyright This work is licensed under a Non Exclusive No Reuse License. Keywords conservation forest regeneration himalayan conifers reproductive biology seed germination Authors Affiliations Abhinay Bhardwaj 0009-0004-8594-3530 [email protected] Forest Research Institute Deemed to be University View all articles by this author Manisha Thapliyal [email protected] ICFRE View all articles by this author Sheeshram Dangwal Forest Research Institute Deemed to be University View all articles by this author Metrics & Citations Metrics Article Usage 360 views 132 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation Abhinay Bhardwaj, Manisha Thapliyal, Sheeshram Dangwal. Seed germination behavior of Tsuga dumosa (D. Don) Eichler (Himalayan Hemlock), a rare gymnosperm of Indian Himalayan Region.. Authorea . 18 July 2025. DOI: https://doi.org/10.22541/au.175283114.49903034/v1 If you have the appropriate software installed, you can download article citation data to the citation manager of your choice. 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