Comparative Study of Cannabis Sativa Ecotypes Found in Three Villages at Lusikisiki, Eastern Cape, South Africa

Authors

  • Azile Dumani

    Department of Biological and Environmental Sciences, Faculty of Natural Science, Walter Sisulu University, Mthatha 5117, South Africa

    Department of Agriculture, Döhne Agricultural Development Institute, Stutterheim 4930, South Africa

  • Tembakazi Theodora Silwana

    Department of Agriculture, Döhne Agricultural Development Institute, Stutterheim 4930, South Africa

  • Ifeanyi Egbichi

    Department of Biological and Environmental Sciences, Faculty of Natural Science, Walter Sisulu University, Mthatha 5117, South Africa

  • Adebola Omowunmi Oyedeji

    Department of Chemical and Physical Sciences, Faculty of Natural Science, Walter Sisulu University, Mthatha 5117, South Africa

  • Babalwa Mpambani

    Department of Agriculture, Döhne Agricultural Development Institute, Stutterheim 4930, South Africa

  • Buyisile Mayekiso

    Department of Botany, University of Fort Hare, Alice 5701, South Africa

  • Hlabana Alfred Seepe

    Department of Agriculture, Land Reform & Rural Development, Quarantine Station, Stellenbosch 7600, South Africa

DOI:

https://doi.org/10.30564/re.v7i5.12235
Received: 25 September 2025 | Revised: 6 November 2025 | Accepted: 14 November 2025 | Published Online: 25 December 2025

Abstract

This study comparatively assesses variation in growth, development, and yield of Cannabis sativa ecotypes from three villages (V1, V2, and V3) in Lusikisiki, Eastern Cape, South Africa. The study was conducted on physiologically mature plants. Six plants per village were selected, and the morphological features, namely, plant height, number of branches, nodes, flowers, plant fresh and dry weights, and flower fresh and dry weights, were determined. One thousand seeds per village were collected, and their weights and moisture levels were determined using a digital scale and a moisture meter. Seeds, leaves, and flowers from each village were fixed and prepared for the determination of anatomical traits, including sizes of the seed, micropyle, and hilum. Trichome types and density on leaves and flowers were determined using a Scanning Electron Microscope (SEM). It was observed that morpho-anatomical traits enable differentiation among ecotypes, influenced by the locally specific environment. Thus, revealing significant variations in vegetative traits (plant height, number of branches, plant fresh and dry weights). V1 recorded the highest vegetative traits, whilst V3 had the highest reproductive and anatomical traits (number of flowers, flower fresh and dry weights, cannabinoid content, trichome density, and seed germinability, aiding features such as the sizes of seed and micropyle). Flowers recorded a higher density of glandular type trichomes than leaves across the villages, although the highest were recorded in V2, followed by V3. In situ conservation of these naturalized cannabis populations through area zoning to ensure genetic preservation for sustainable product supply and economic growth is recommended.

Keywords:

Cannabis; Ecotype; Variation; Morphological Traits; Anatomy; Preservation; Ecology

References

[1] Panahi, S., KhandanMirkohi, A., Taylor, G., et al., 2024. Characterizing Morphological Properties of Select Populations of Iranian Fiber Cannabis Cannabis sativa L.). International Journal of Horticultural Science and Technology. 11(1). DOI: https://doi.org/10.22059/ijhst.2023.349507.591

[2] Andre, C.M., Hausman, J.-F., Guerriero, G., 2016. Cannabis sativa: The Plant of the Thousand and One Molecules. Frontiers in Plant Science. 7. DOI: https://doi.org/10.3389/fpls.2016.00019

[3] Shahzad, A., 2012. Hemp Fiber and Its Composites – a Review. Journal of Composite Materials. 46(8), 973–986. DOI: https://doi.org/10.1177/0021998311413623

[4] Simiyu, D.C., Jang, J.H., Lee, O.R., 2022. Understanding Cannabis sativa L.: Current Status of Propagation, Use, Legalization, and Haploid-Inducer-Mediated Genetic Engineering. Plants. 11(9), 1236. DOI: https://doi.org/10.3390/plants11091236

[5] Ren, G., Zhang, X., Li, Y., et al., 2021. Large-Scale Whole-Genome Resequencing Unravels the Domestication History of Cannabis Sativa. Science Advances. 7(29), eabg2286. DOI: https://doi.org/10.1126/sciadv.abg2286

[6] Zhang, Q., Chen, X., Guo, H., et al., 2018. Latitudinal Adaptation and Genetic Insights Into the Origins of Cannabis sativa L. Frontiers in Plant Science. 9, 1876. DOI: https://doi.org/10.3389/fpls.2018.01876

[7] Hesami, M., Pepe, M., Jones, A.M.P., 2023. Morphological Characterization of Cannabis sativa L. Throughout Its Complete Life Cycle. Plants. 12(20), 3646. DOI: https://doi.org/10.3390/plants12203646

[8] Isahq, M.S., Afridi, M.S., Ali, J., et al., 2015. Proximate Composition, Phytochemical Screening, GC-MS Studies of Biologically Active Cannabinoids and Antimicrobial Activities of Cannabis Indica. Asian Pacific Journal of Tropical Disease. 5(11), 897–902. DOI: https://doi.org/10.1016/S2222-1808(15)60953-7

[9] Gülck, T., Booth, J.K., Carvalho, Â., et al., 2020. Synthetic Biology of Cannabinoids and Cannabinoid Glucosides in Nicotiana benthamiana and Saccharomyces cerevisiae. Journal of Natural Products. 83(10), 2877–2893. DOI: https://doi.org/10.1021/acs.jnatprod.0c00241

[10] Seddon, T., Floodgate, W., 2020. A Global Review of Cannabis Regulation Models. In: Regulating Cannabis. Springer International Publishing: Cham, Switzerland. pp. 15–83. DOI: https://doi.org/10.1007/978-3-030-52927-7_2

[11] Duvall, C.S., 2019. A Brief Agricultural History of Cannabis in Africa, from Prehistory to Canna-Colony. EchoGéo. 48. DOI: https://doi.org/10.4000/echogeo.17599

[12] Kitchen, C., Kabba, J.A., Fang, Y., 2022. Status and Impacts of Recreational and Medicinal Cannabis Policies in Africa: A Systematic Review and Thematic Analysis of Published and “Gray” Literature. Cannabis and Cannabinoid Research. 7(3), 239–261. DOI: https://doi.org/10.1089/can.2021.0110

[13] Adhikary, D., Kulkarni, M., El-Mezawy, A., et al., 2021. Medical Cannabis and Industrial Hemp Tissue Culture: Present Status and Future Potential. Frontiers in Plant Science. 12, 627240. DOI: https://doi.org/10.3389/fpls.2021.627240

[14] Fiorentino, N., Formisano, C., Delfine, S., et al., 2024. Editorial: Environmental and Agronomic Factors Affecting the Chemical Composition and Biological Activity of Cannabis Extracts. Frontiers in Plant Science. 15, 1407262. DOI: https://doi.org/10.3389/fpls.2024.1407262

[15] Kim, S., 2024. Genetic and Environmental Factors Shaping Cannabis Phenotypes: A Study on Temperature Effects and Genetic Regulation of Anthocyanin Accumulation in Cannabis sativa [PhD Thesis]. University of Wisconsin-Madison: Madison, WI, USA. Available from: http://digital.library.wisc.edu/1793/85243.

[16] Payment, J., Cvetkovska, M., 2023. The Responses of Cannabis Sativa to Environmental Stress: A Balancing Act. Botany. 101(8), 318–332. DOI: https://doi.org/10.1139/cjb-2023-0056

[17] Ahsan, S.M., Injamum-Ul-Hoque, Md., Shaffique, S., et al., 2024. Illuminating Cannabis Sativa L.: The Power of Light in Enhancing C. Sativa Growth and Secondary Metabolite Production. Plants. 13(19), 2774. DOI: https://doi.org/10.3390/plants13192774

[18] Razzaq, K., Du, J., 2025. Phytohormonal Regulation of Plant Development in Response to Fluctuating Light Conditions. Journal of Plant Growth Regulation. 44(5), 1903–1936. DOI: https://doi.org/10.1007/s00344-024-11568-5

[19] Li, M., Roman, M., Yuan, J., et al., 2023. Varying Light Intensity Can Alter Metabolic Profile and Cannabispiradienone Content of Industrial Hemp. Industrial Crops and Products. 202, 117031. DOI: https://doi.org/10.1016/j.indcrop.2023.117031

[20] Langa, S., Magwaza, L.S., Mditshwa, A., et al., 2024. Temperature Effects on Seed Germination and Seedling Biochemical Profile of Cannabis Landraces. International Journal of Plant Biology. 15(4), 1032–1053. DOI: https://doi.org/10.3390/ijpb15040073

[21] Rehman, A., Khan, I., Farooq, M., 2024. Secondary Metabolites Mediated Reproductive Tolerance Under Heat Stress in Plants. Journal of Plant Growth Regulation. 43(9), 2993–3011. DOI: https://doi.org/10.1007/s00344-023-11161-2

[22] Archer, R., 2025. Optimizing CBD Hemp Production in Cold Climates: Accession, High-Tunnel, and Frost Effects on Yield and Cannabinoid Profiles [Master's thesis]. North Dakota State University: Fargo, ND, USA.

[23] Ogwu, M.C., Izah, S.C., Joshua, M.T., 2025. Ecological and Environmental Determinants of Phytochemical Variability in Forest Trees. Phytochemistry Reviews. DOI: https://doi.org/10.1007/s11101-025-10066-0

[24] Da Cunha Leme Filho, J.F., Schuchman, S., De Sarandy Raposo, R., et al., 2025. The Role of Jasmonates in Modulating Growth, Trichome Density, and Cannabinoid Accumulation in Cannabis sativa L. International Journal of Plant Biology. 16(2), 68. DOI: https://doi.org/10.3390/ijpb16020068

[25] Scharwies, J.D., Dinneny, J.R., 2019. Water Transport, Perception, and Response in Plants. Journal of Plant Research. 132(3), 311–324. DOI: https://doi.org/10.1007/s10265-019-01089-8

[26] Zhao, W., Liu, L., Shen, Q., et al., 2020. Effects of Water Stress on Photosynthesis, Yield, and Water Use Efficiency in Winter Wheat. Water. 12(8), 2127. DOI: https://doi.org/10.3390/w12082127

[27] Cao, W., Sun, H., Shao, C., et al., 2025. Progress in the Study of Plant Nitrogen and Potassium Nutrition and Their Interaction Mechanisms. Horticulturae. 11(8), 930. DOI: https://doi.org/10.3390/horticulturae11080930

[28] Song, C., Saloner, A., Fait, A., et al., 2023. Nitrogen Deficiency Stimulates Cannabinoid Biosynthesis in Medical Cannabis Plants by Inducing a Metabolic Shift Towards Production of Low-N Metabolites. Industrial Crops and Products. 202, 116969. DOI: https://doi.org/10.1016/j.indcrop.2023.116969

[29] Hesami, M., 2024. Morphological and Genetic Dynamics in Cannabis sativa L.: Transcriptomic Insights into Leaf Morphogenesis, Phase Transition, and Somatic Embryogenesis [PhD Thesis]. University of Guelph: Guelph, ON, Canada. Available from: https://hdl.handle.net/10214/28107.

[30] Kovalchuk, I., Pellino, M., Rigault, P., et al., 2020. The Genomics of Cannabis and Its Close Relatives. Annual Review of Plant Biology. 71(1), 713–739. DOI: https://doi.org/10.1146/annurev-arplant-081519-040203

[31] Caplan, D.M., 2018. Propagation and Root Zone Management For Controlled Environment Cannabis Production [PhD Thesis]. University of Guelph: Guelph, ON, Canada. Available from: http://hdl.handle.net/10214/14249

[32] Carlson, C.H., Stack, G.M., Jiang, Y., et al., 2021. Morphometric Relationships and Their Contribution to Biomass and Cannabinoid Yield in Hybrids of Hemp ( Cannabis Sativa ). Journal of Experimental Botany. 72(22), 7694–7709. DOI: https://doi.org/10.1093/jxb/erab346

[33] Bellon, M.R., Dulloo, E., Sardos, J., et al., 2017. In Situ Conservation—Harnessing Natural and Human‐Derived Evolutionary Forces to Ensure Future Crop Adaptation. Evolutionary Applications. 10(10), 965–977. DOI: https://doi.org/10.1111/eva.12521

[34] Jin, D., Dai, K., Xie, Z., et al., 2020. Secondary Metabolites Profiled in Cannabis Inflorescences, Leaves, Stem Barks, and Roots for Medicinal Purposes. Scientific Reports. 10(1), 3309. DOI: https://doi.org/10.1038/s41598-020-60172-6

[35] Shams, R., Azizi, A., Hamzei, J., et al., 2020. Genetic Structure and Diversity of Iranian Cannabis Populations Based on Phytochemical, Agro-Morphological and Molecular Markers. Industrial Crops and Products. 158, 112950. DOI: https://doi.org/10.1016/j.indcrop.2020.112950

[36] Conneely, L.J., Mauleon, R., Mieog, J., et al., 2021. Characterization of the Cannabis Sativa Glandular Trichome Proteome. PLOS ONE. 16(4), e0242633. DOI: https://doi.org/10.1371/journal.pone.0242633

[37] Del Rosario-Makridis, G., 2023. Molecular Regulation of Glandular Trichome Initiation and Morphogenesis in Cannabis sativa L [PhD Thesis]. La Trobe University: Melbourne, Australia. DOI: https://doi.org/10.26181/25744944.v1

[38] Tanney, C.A.S., Backer, R., Geitmann, A., et al., 2021. Cannabis Glandular Trichomes: A Cellular Metabolite Factory. Frontiers in Plant Science. 12, 721986. DOI: https://doi.org/10.3389/fpls.2021.721986

[39] Shukla, P., Prasad, A., Chawda, K., et al, 2024. Glandular trichomes: Bio-cell factories of plant secondary metabolites. In: In Vitro Propagation and Secondary Metabolite Production from Medicinal Plants: Current Trends (Part 1). Bentham Science Publishers: Singapore. pp. 91–119.

[40] Lawrence, R.G., 2019. Pot in pans: A history of eating cannabis. Rowman & Littlefield.

[41] Żuk-Gołaszewska, K., Żuk-Gołaszewska, K., Gołaszewski, J., 2018. Cannabis Sativa L. – Cultivation and Quality of Raw Material. Journal of Elementology. (3/2018). DOI: https://doi.org/10.5601/jelem.2017.22.3.1500

[42] Ibarra-Lecue, I., Pilar-Cuéllar, F., Muguruza, C., et al., 2018. The Endocannabinoid System in Mental Disorders: Evidence from Human Brain Studies. Biochemical Pharmacology. 157, 97–107. DOI: https://doi.org/10.1016/j.bcp.2018.07.009

[43] Mostafaei Dehnavi, M., Ebadi, A., Peirovi, A., et al., 2022. THC and CBD Fingerprinting of an Elite Cannabis Collection from Iran: Quantifying Diversity to Underpin Future Cannabis Breeding. Plants. 11(1), 129. DOI: https://doi.org/10.3390/plants11010129

[44] Datwyler, S.L., Weiblen, G.D., 2006. Genetic Variation in Hemp and Marijuana ( Cannabis sativa L.) According to Amplified Fragment Length Polymorphisms. Journal of Forensic Sciences. 51(2), 371–375. DOI: https://doi.org/10.1111/j.1556-4029.2006.00061.x

[45] Potter, D.J., 2014. A Review of the Cultivation and Processing of Cannabis ( Cannabis Sativa L.) for Production of Prescription Medicines in the UK. Drug Testing and Analysis. 6(1–2), 31–38. DOI: https://doi.org/10.1002/dta.1531

[46] Taura, F., Sirikantaramas, S., Shoyama, et al., 2007. Cannabidiolic‐Acid Synthase, the Chemotype‐Determining Enzyme in the Fiber‐Type Cannabis Sativa. FEBS Letters. 581(16), 2929–2934. DOI: https://doi.org/10.1016/j.febslet.2007.05.043

[47] Vlad, R.A., Hancu, G., Ciurba, A., et al., 2020. Cannabidiol-Therapeutic and Legal Aspects. Die Pharmazie-An International Journal of Pharmaceutical Sciences. 75(10), 463–469. Available from: https://www.ingentaconnect.com/content/govi/pharmaz/2020/00000075/00000010/art00002;jsessionid=10vws302age0n.x-ic-live-02

[48] Zendulka, O., Dovrtělová, G., Nosková, K., et al., 2016. Cannabinoids and Cytochrome P450 Interactions. Current Drug Metabolism. 17(3), 206–226. DOI: https://doi.org/10.2174/1389200217666151210142051

[49] Murovec, J., Eržen, J.J., Flajšman, M., et al., 2022. Analysis of Morphological Traits, Cannabinoid Profiles, THCAS Gene Sequences, and Photosynthesis in Wide and Narrow Leaflet High-Cannabidiol Breeding Populations of Medical Cannabis. Frontiers in Plant Science. 13, 786161. DOI: https://doi.org/10.3389/fpls.2022.786161

[50] Owen-Smith, N., 2021. Only in Africa: The Ecology of Human Evolution, 1st ed. Cambridge University Press: Cambridge, UK. DOI: https://doi.org/10.1017/9781108961646

[51] Maponya, P., Madakadze, C., Mbili, N., et al., 2023. Potential Constraint of Rainfall Availability on the Establishment and Expansion of Agroforestry in the Joe Gqabi, Alfred Nzo and OR Tambo Districts, Eastern Cape in South Africa. In: Agricultural Bioeconomy. Elsevier: London, UK. pp. 121–141. DOI: https://doi.org/10.1016/B978-0-323-90569-5.00009-3

[52] Mucina, L., Rutherford, M.C., Powrie, L.W., 2006. Vegetation Atlas of South Africa, Lesotho and Swaziland. South African National Biodiversity Institute: Pretoria, South Africa. pp. 748–789.

[53] Cheteni, P., Khamfula, Y., Mah, G., 2020. Exploring Food Security and Household Dietary Diversity in the Eastern Cape Province, South Africa. Sustainability. 12(5), 1851. DOI: https://doi.org/10.3390/su12051851

[54] Nkamisa, M., Ndhleve, S., Nakin, M.D.V., et al., 2022. Analysis of Trends, Recurrences, Severity and Frequency of Droughts Using Standardised Precipitation Index: Case of OR Tambo District Municipality, Eastern Cape, South Africa. Jàmbá: Journal of Disaster Risk Studies. 14(1). DOI: https://doi.org/10.4102/jamba.v14i1.1147

[55] Tipple, B.J., Hambach, B., Barnette, J.E., et al., 2016. The Influences of Cultivation Setting on Inflorescence Lipid Distributions, Concentrations, and Carbon Isotope Ratios of Cannabis Sp. Forensic Science International. 262, 233–241. DOI: https://doi.org/10.1016/j.forsciint.2016.03.029

[56] Demirel, C., Kabutey, A., Herák, D., et al., 2021. Optimizing Uniaxial Oil Extraction of Bulk Rapeseeds: Spectrophotometric and Chemical Analyses of the Extracted Oil under Pretreatment Temperatures and Heating Intervals. Processes. 9(10), 1755. DOI: https://doi.org/10.3390/pr9101755

[57] Olascuaga-Castillo, K.A., Castillo-Medina, O., Villacorta-Zavaleta, M., et al., 2024. Extraction of Essential Oils by Hydrodistillation of Four Aromatic Species: Conditioning, Extraction Conditions, Yield and Chemical Composition. Scientia Agropecuaria. 15(3), 385–408. DOI: https://doi.org/10.17268/sci.agropecu.2024.029

[58] Alzarieni, K.Z., Zhang, Y., Niyonsaba, E., et al., 2021. Determination of the Chemical Compositions of Condensate-like Oils with Different API Gravities by Using the Distillation, Precipitation, Fractionation Mass Spectrometry (DPF MS) Method. Energy & Fuels. 35(10), 8646–8656. DOI: https://doi.org/10.1021/acs.energyfuels.0c04286

[59] Aljamali, N.M., Salih, N.S., 2021. Review on Chemical Separation Of Crude Oil And Analysis Of Its Components. Journal of Petroleum Engineering & Technology, 11(2), 35–49. DOI: https://doi.org/10.37591/JoPET.

[60] Turner, C., Cheng, P., Lewis, G., et al., 1979. Constituents of Cannabis sativa. Planta Medica. 37(11), 217–225. DOI: https://doi.org/10.1055/s-0028-1097331

[61] Nor Asikin, N.A.N., Abu Bakar, N.S., Muhammad Nor, S.M., et al., 2021. Density And Structure Of Leaf Trichomes In Capsicum Annuum and Capsicum Frutescens. Universiti Malaysia Terengganu Journal of Undergraduate Research. 3(3), 81–86. DOI: https://doi.org/10.46754/umtjur.v3i3.220

[62] Lemus-Barrios, H., Barrios, D., García-Beltrán, J.A., et al., 2021. Taxonomic Implications of Seed Morphology in Melocactus (Cactaceae) from Cuba. Willdenowia. 51(1). DOI: https://doi.org/10.3372/wi.51.51108

[63] Talip, N., Abdul Rahman, M.R. and Ahmad Juhari, M.A.A., 2019. Plant Anatomy and Microscopy. Universiti Kebangsaan Malaysia Press: Bangi, Malaysia. (in Malaysian)

[64] Abdelhameed, A., Amer, W., Hassan, W., et al., 2020. Auto-Taxonomy of Brassica Tournefortii Gouan. (Brassicaceae) in Egypt. Bangladesh Journal of Plant Taxonomy. 27(2), 233–250. DOI: https://doi.org/10.3329/bjpt.v27i2.50664

[65] Fisher, J.B., Malhi, Y., Torres, I.C., et al., 2013. Nutrient Limitation in Rainforests and Cloud Forests Along a 3,000-M Elevation Gradient in the Peruvian Andes. Oecologia. 172(3), 889–902. DOI: https://doi.org/10.1007/s00442-012-2522-6

[66] He, X., Hou, E., Liu, Y., et al., 2016. Altitudinal Patterns and Controls of Plant and Soil Nutrient Concentrations and Stoichiometry in Subtropical China. Scientific Reports. 6(1), 24261. DOI: https://doi.org/10.1038/srep24261

[67] Zhu, D., Zhou, Y., Peng, S., et al., 2024. Impacts of Altitude on Plant Green Leaf, Fresh Litter, and Soil Stoichiometry in Subtropical Forests. Frontiers in Forests and Global Change. 7, 1331623. DOI: https://doi.org/10.3389/ffgc.2024.1331623

[68] Singh, S.., 2018. Understanding the Role Of Slope Aspect In Shaping The Vegetation Attributes And Soil Properties In Montane Ecosystems. Tropical Ecology. 59(3), 417–430. Available from: https://www.researchgate.net/publication/341354955_Understanding_the_role_of_slope_aspect_in_shaping_the_vegetation_attributes_and_soil_properties_in_Montane_ecosystems

[69] Filimonenko, E., Kuzyakov, Y., 2025. Activation Energy of Organic Matter Decomposition in Soil and Consequences of Global Warming. Global Change Biology. 31(9), e70472. DOI: https://doi.org/10.1111/gcb.70472

[70] Xue, J., 2018. Plant Traits and Biomass Allocation of Gentiana Hexaphylla on Different Slope Aspects at the Eastern Margin of Qinghai-Tibet Plateau. Applied Ecology and Environmental Research. 16(2), 1835–1853. DOI: https://doi.org/10.15666/aeer/1602_18351853

[71] Grassein, F., Till-Bottraud, I., Lavorel, S., 2010. Plant Resource-Use Strategies: The Importance of Phenotypic Plasticity in Response to a Productivity Gradient for Two Subalpine Species. Annals of Botany. 106(4), 637–645. DOI: https://doi.org/10.1093/aob/mcq154

[72] Ghosh, B.N., Sharma, N.K., Alam, N.M., et al., 2014. Elevation, Slope Aspect and Integrated Nutrient Management Effects on Crop Productivity and Soil Quality in North-West Himalayas, India. Journal of Mountain Science. 11(5), 1208–1217. DOI: https://doi.org/10.1007/s11629-013-2674-9

[73] Körner, C., 1998. A Re-Assessment of High Elevation Treeline Positions and Their Explanation. Oecologia. 115(4), 445–459. DOI: https://doi.org/10.1007/s004420050540

[74] Méndez, M., Traveset, A., 2003. Sexual Allocation in Single-Flowered Hermaphroditic Individuals in Relation to Plant and Flower Size. Oecologia. 137(1), 69–75. DOI: https://doi.org/10.1007/s00442-003-1319-z

[75] He, J., Xue, J., Gao, J., et al., 2017. Adaptations of the Floral Characteristics and Biomass Allocation Patterns of Gentiana Hexaphylla to the Altitudinal Gradient of the Eastern Qinghai-Tibet Plateau. Journal of Mountain Science. 14(8), 1563–1576. DOI: https://doi.org/10.1007/s11629-017-4424-x

[76] Li, D., Wang, X., Zhang, X., et al., 2016. The Genetic Architecture of Leaf Number and Its Genetic Relationship to Flowering Time in Maize. New Phytologist. 210(1), 256–268. DOI: https://doi.org/10.1111/nph.13765

[77] Goodwillie, C., Sargent, R.D., Eckert, C.G., et al., 2010. Correlated Evolution of Mating System and Floral Display Traits in Flowering Plants and Its Implications for the Distribution of Mating System Variation. New Phytologist. 185(1), 311–321. DOI: https://doi.org/10.1111/j.1469-8137.2009.03043.x

[78] Rodriguez-Morrison, V., Llewellyn, D., Zheng, Y., 2021. Cannabis Yield, Potency, and Leaf Photosynthesis Respond Differently to Increasing Light Levels in an Indoor Environment. Frontiers in Plant Science. 12, 646020. DOI: https://doi.org/10.3389/fpls.2021.646020

[79] Poorter, H., Niklas, K.J., Reich, P.B., et al., 2012. Biomass Allocation to Leaves, Stems and Roots: Meta‐Analyses of Interspecific Variation and Environmental Control. New Phytologist. 193(1), 30–50. DOI: https://doi.org/10.1111/j.1469-8137.2011.03952.x

[80] Eichhorn Bilodeau, S., Wu, B.-S., Rufyikiri, A.-S., et al., 2019. An Update on Plant Photobiology and Implications for Cannabis Production. Frontiers in Plant Science. 10, 296. DOI: https://doi.org/10.3389/fpls.2019.00296

[81] McPartland, J.M., Guy, G.W., Hegman, W., 2018. Cannabis Is Indigenous to Europe and Cultivation Began During the Copper or Bronze Age: A Probabilistic Synthesis of Fossil Pollen Studies. Vegetation History and Archaeobotany. 27(4), 635–648. DOI: https://doi.org/10.1007/s00334-018-0678-7

[82] Dang, M., Arachchige, N.M., Campbell, L.G., 2022. Optimizing Photoperiod Switch to Maximize Floral Biomass and Cannabinoid Yield in Cannabis sativa L.: A Meta-Analytic Quantile Regression Approach. Frontiers in Plant Science. 12, 797425. DOI: https://doi.org/10.3389/fpls.2021.797425

[83] Cheng, X., Zhao, K., He, X., et al., 2023. Differential Protein Response to Different Light Quality Conditions of Industrial Hemp Cultivation Based on DIA Technology. Industrial Crops and Products. 197, 116650. DOI: https://doi.org/10.1016/j.indcrop.2023.116650

[84] Abdollahi, M., Sefidkon, F., Calagari, M., et al., 2020. A Comparative Study of Seed Yield and Oil Composition of Four Cultivars of Hemp (Cannabis Sativa L.) Grown from Three Regions in Northern Iran. Industrial Crops and Products. 152, 112397. DOI: https://doi.org/10.1016/j.indcrop.2020.112397

[85] Petit, J., Salentijn, E.M.J., Paulo, M.-J., et al., 2020. Genetic Variability of Morphological, Flowering, and Biomass Quality Traits in Hemp (Cannabis sativa L.). Frontiers in Plant Science. 11, 102. DOI: https://doi.org/10.3389/fpls.2020.00102

[86] Baldini, M., Ferfuia, C., Piani, B., et al., 2018. The Performance and Potentiality of Monoecious Hemp (Cannabis sativa L.) Cultivars as a Multipurpose Crop. Agronomy. 8(9), 162. DOI: https://doi.org/10.3390/agronomy8090162

[87] Dilena, E., Close, D.C., Hunt, I., et al., 2023. Investigating How Nitrogen Nutrition and Pruning Impacts on CBD and THC Concentration and Plant Biomass of Cannabis Sativa. Scientific Reports. 13(1), 19533. DOI: https://doi.org/10.1038/s41598-023-46369-5

[88] Nabila, C.I.B., Ramona, Y., Wirasuta, I.M.A.G., et al., 2025. Factors Affecting Cannabidiol and Tetrahydrocannabinol Production in Cannabis: Internal Mechanisms and Environmental Factors – A Systematic Review. Journal of Pharmacy & Pharmacognosy Research. 13(4), 1178–1190. DOI: https://doi.org/10.56499/jppres24.2184_13.4.1178

[89] Trancoso, I., De Souza, G.A.R., Dos Santos, P.R., et al., 2022. Cannabis sativa L.: Crop Management and Abiotic Factors That Affect Phytocannabinoid Production. Agronomy. 12(7), 1492. DOI: https://doi.org/10.3390/agronomy12071492

[90] Alsaleh, A., Yılmaz, G., 2025. Exploring Cannabidiol Variations, Investigation of Genetic Diversity, Population Structure and Unveiling Male-Specific Genetic Marker in Industrial Hemp (Cannabis Sativa L.). Genetic Resources and Crop Evolution. 72(1), 797–814. DOI: https://doi.org/10.1007/s10722-024-02015-1

[91] Campbell, B.J., Berrada, A.F., Hudalla, C., et al., 2019. Genotype × Environment Interactions of Industrial Hemp Cultivars Highlight Diverse Responses to Environmental Factors. Agrosystems, Geosciences & Environment. 2(1), 1–11. DOI: https://doi.org/10.2134/age2018.11.0057

[92] Khan, A., Kanwal, F., Ullah, S., et al., 2025. Plant Secondary Metabolites—Central Regulators Against Abiotic and Biotic Stresses. Metabolites. 15(4), 276. DOI: https://doi.org/10.3390/metabo15040276

[93] Ostovary, I., 2016. The Influence Of Nurse Shrubs On Growth And Establishment Of Olea Europaea Subsp. Cuspidata [PhD Thesis]. University of Natural Resources and Life Sciences Vienna: Vienna, Austria. Available from: https://www.semanticscholar.org/paper/The-influence-of-nurse-shrubs-on-growth-and-of-Olea-Godbold-Godbold/959aa1339bdf99093d0d72c5590cee1341bc328a

[94] Rashid, N., Zafar, M., Ahmad, M., et al., 2018. Intraspecific Variation in Seed Morphology of Tribe Vicieae (Papilionoidae) Using Scanning Electron Microscopy Techniques. Microscopy Research and Technique. 81(3), 298–307. DOI: https://doi.org/10.1002/jemt.22979

[95] Davitashvili, N., Karrer, G., 2010. Taxonomic Importance of Seed Morphology in Gentiana (Gentianaceae): Taxonomic Importance Of Seed Morphology In Gentiana. Botanical Journal of the Linnean Society. 162(1), 101–115. DOI: https://doi.org/10.1111/j.1095-8339.2009.01020.x

[96] Rashid, N., Zafar, M., Ahmad, M., et al., 2021. Seed morphology: An addition to the taxonomy of Astragaleae and Trifolieae (Leguminosae: Papilionoidae) from Pakistan. Microscopy Research and Technique. 84(5), 1053–1062. DOI: https://doi.org/10.1002/jemt.23666

[97] Moon, Y., Cha, Y., Lee, J., et al., 2020. Investigation of Suitable Seed Sizes, Segregation of Ripe Seeds, and Improved Germination Rate for the Commercial Production of Hemp Sprouts ( Cannabis Sativa L.). Journal of the Science of Food and Agriculture. 100(7), 2819–2827. DOI: https://doi.org/10.1002/jsfa.10294

[98] Langa, S., Magwaza, L.S., Mditshwa, A., et al., 2024. Characterization of Cannabis Varieties and the Intrinsic and Extrinsic Factors Affecting Cannabis Germination and Seedling Establishment: A Descriptive Review. Industrial Crops and Products. 208, 117861. DOI: https://doi.org/10.1016/j.indcrop.2023.117861

[99] Waheed, A., Ahmad, M., Ghufran, M.A., et al., 2021. Implication of Scanning Electron Microscopy in the Seed Morphology with Special Reference to Three Subfamilies of Fabaceae. Microscopy Research and Technique. 84(9), 2176–2185. DOI: https://doi.org/10.1002/jemt.23772

[100] Kaya, A., Ünal, M., Özgökçe, F., et al., 2011. Fruit and Seed Morphology of Six Species Previously Placed in Malcolmia (Brassicaceae) in Turkey and Their Taxonomic Value. Turkish Journal of Botany. DOI: https://doi.org/10.3906/bot-1010-99

[101] Stoilkovska Gjorgievska, V., Geskovski, N., Makreski, P., et al., 2024. Differentiation of Cannabis Seeds Employing Digital Morphological Screening and Infrared Spectroscopy Coupled with Multivariate Modeling. Industrial Crops and Products. 211, 118184. DOI: https://doi.org/10.1016/j.indcrop.2024.118184

[102] ElSohly, M.A., Radwan, M.M., Gul, W., et al., 2017. Phytochemistry of Cannabis sativa L. In: Kinghorn, A.D., Falk, H., Gibbons, S., et al. (Eds.). Phytocannabinoids, Progress in the Chemistry of Organic Natural Products. Springer International Publishing, Cham, Switzerland. pp. 1–36. DOI: https://doi.org/10.1007/978-3-319-45541-9_1

[103] Zhao, X., Liu, Y., Li, J., et al., 2023. Numerical Analyses of Seed Morphology and Its Taxonomic Significance in the Genus Oxytropis DC. (Fabaceae) from Northwestern China. PhytoKeys. 222, 49–67. DOI: https://doi.org/10.3897/phytokeys.222.96990

[104] Boesewinkel, F.D., Bouman, F., 2017. The Seed: Structure And Function. In: Seed development and germination. Routledge: London, UK. pp. 1–24.

[105] Coffigniez, F., Briffaz, A., Mestres, C., et al., 2019. Impact of Soaking Process on the Microstructure of Cowpea Seeds in Relation to Solid Losses and Water Absorption. Food Research International. 119, 268–275. DOI: https://doi.org/10.1016/j.foodres.2019.02.010

[106] Zewdie, T., Welka, K., 2015. Effect of Micropyle Orientation on Germination of Millettia Ferruginea and Delonix Regia. Ecological Processes. 4(1), 12. DOI: https://doi.org/10.1186/s13717-015-0038-9

[107] Doyle, J.A., 2017. Phylogenetic Analyses and Morphological Innovations in Land Plants. In: Roberts, J.A. (Ed.). Annual Plant Reviews Online. Wiley: London, UK. pp. 1–50. DOI: https://doi.org/10.1002/9781119312994.apr0486

[108] Taiz, L., Taiz, L., 2017. Flora Unveiled: The Discovery And Denial Of Sex In Plants. Oxford University Press: Oxford, UK.

[109] Zhao, Q., Shi, X., Yan, L., et al., 2021. Characterization of the Common Genetic Basis Underlying Seed Hilum Size, Yield, and Quality Traits in Soybean. Frontiers in Plant Science. 12, 610214. DOI: https://doi.org/10.3389/fpls.2021.610214

[110] Sutton, D.B., Punja, Z.K., Hamarneh, G., 2023. Characterization of Trichome Phenotypes to Assess Maturation and Flower Development in Cannabis Sativa L. (cannabis) by Automatic Trichome Gland Analysis. Smart Agricultural Technology. 3, 100111. DOI: https://doi.org/10.1016/j.atech.2022.100111

[111] Livingston, S.J., Quilichini, T.D., Booth, J.K., et al., 2020. Cannabis Glandular Trichomes Alter Morphology and Metabolite Content During Flower Maturation. The Plant Journal. 101(1), 37–56. DOI: https://doi.org/10.1111/tpj.14516

[112] Ahrens, A., Llewellyn, D., Zheng, Y., 2024. Longer Photoperiod Substantially Increases Indoor-Grown Cannabis’ Yield and Quality: A Study of Two High-THC Cultivars Grown under 12 h vs. 13 h Days. Plants. 13(3), 433. DOI: https://doi.org/10.3390/plants13030433

[113] Punja, Z.K., Sutton, D.B., Kim, T., 2023. Glandular Trichome Development, Morphology, and Maturation Are Influenced by Plant Age and Genotype in High THC-Containing Cannabis (Cannabis Sativa L.) Inflorescences. Journal of Cannabis Research. 5(1), 12. DOI: https://doi.org/10.1186/s42238-023-00178-9

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Dumani, A., Silwana, T. T., Egbichi, I., Oyedeji, A. O., Mpambani, B., Mayekiso, B., & Seepe, H. A. (2025). Comparative Study of Cannabis Sativa Ecotypes Found in Three Villages at Lusikisiki, Eastern Cape, South Africa. Research in Ecology, 7(5), 336–356. https://doi.org/10.30564/re.v7i5.12235