ElSohly, M. A. & Slade, D. Chemical constituents of marijuana: The complex mixture of natural cannabinoids. Life Sci. 78, 539–548 (2005).
Radwan, M. et al. Isolation and characterization of new Cannabis constituents from a high potency variety. Planta Med. 74, 267–272 (2008).
Poklis, J. L., Thompson, C. C., Long, K. A., Lichtman, A. H. & Poklis, A. Disposition of cannabichromene, cannabidiol, and 9-tetrahydrocannabinol and its metabolites in mouse brain following marijuana inhalation determined by high-performance liquid chromatography-tandem mass spectrometry. J. Anal. Toxicol. 34, 516–520 (2010).
Mehmedic, Z. et al. Potency trends of Δ9-THC and other cannabinoids in confiscated Cannabis preparations from 1993 to 2008*. J. Forensic Sci. 55, 1209–1217 (2010).
Fellermeier, M., Eisenreich, W., Bacher, A. & Zenk, M. H. Biosynthesis of cannabinoids. Eur. J. Biochem. 268, 1596–1604 (2001).
Johnson, M. S. & Wallace, J. G. Genomic and chemical diversity of commercially available High-CBD industrial hemp accessions. Front. Genet. 12 (2021).
de Meijer, E. P. M. et al. The inheritance of chemical phenotype in Cannabis sativa L. Genetics 163, 335–346 (2003).
Staginnus, C., Zörntlein, S. & de Meijer, E. A PCR marker linked to a THCA synthase polymorphism is a reliable tool to discriminate potentially THC-Rich plants of Cannabis sativa L. J. Forensic Sci. 59, 919–926 (2014).
Grassa, C. J. et al. A new Cannabis genome assembly associates elevated Cannabidiol (CBD) with hemp introgressed into marijuana. New Phytol. 230, 1665–1679 (2021).
Grassi, G. & McPartland, J. M. Chemical and morphological phenotypes in breeding of Cannabis sativa L. In Cannabis sativa L. – Botany and Biotechnology 137–160 (Springer International Publishing, 2017).
Weiblen, G. D. et al. Gene duplication and divergence affecting drug content in Cannabis sativa. New Phytol. 208, 1241–1250 (2015).
van Velzen, R. & Schranz, M. E. Origin and evolution of the cannabinoid oxidocyclase gene family. Genome Biol. Evol. 13 (2021).
US Congress. Agricultural Act of 2014 113–179 (Senate and House of Representatives of the United States of America, 2014).
US Congress. Agriculture Improvement Act of 2018 115–334 (Senate and House of Representatives of the United States of America, 2018).
Fike, J. I. Industrial hemp: Renewed opportunities for an ancient crop. CRC Crit. Rev. Plant. Sci. 35, 406–424 (2016).
Schoenrock, R. E. Hemp in Minnesota During the Wartime Emergency (Mankato State College, 1966).
Robinson, R. The Great Book of Hemp: The Complete Guide to the Environmental, Commercial and Medicinal Uses of the World’s Most Extraordinary Plant Vols. 978-089281541–8 (1996).
Wilsie, C. P. & Black, C. A. Hemp production experiments cultural practices and soil requirements. Iowa State Univ. Digit. Repos. 3 (1944).
Haney, A. & Kutscheid, B. B. An ecological study of naturalized hemp (Cannabis sativa L.) in East-Central Illinois. Am. Midl. Nat. 93, 1 (1975).
Lyster, H. & Dewey Hemp: USDA Yearbook of Agriculture (1914).
Stack, G. M. et al. Comparison of recombination rate, reference bias, and unique pangenomic haplotypes in Cannabis sativa using seven de Novo genome assemblies. Int. J. Mol. Sci. 26, 1165 (2025).
Lynch, R. C. et al. Domesticated cannabinoid synthases amid a wild mosaic cannabis pangenome. https://doi.org/10.1101/2024.05.21.595196 (2024).
Faux, A. M., Draye, X., Flamand, M. C., Occre, A. & Bertin, P. Identification of QTLs for sex expression in dioecious and monoecious hemp (Cannabis sativa L). Euphytica 209, 357–376 (2016).
Petit, J. et al. Elucidating the genetic architecture of fiber quality in hemp (Cannabis sativa L.) using a genome-wide association study. Front. Genet. 11 (2020).
de Ronne, M., Lapierre, É. & Torkamaneh, D. Genetic insights into agronomic and morphological traits of drug-type cannabis revealed by genome-wide association studies. Sci. Rep. 14, 9162 (2024).
GRIN-Global. U.S. National Plant Germplasm System. GRIN-Glob. Web version 11040. GRIN-Global. https://npgsweb.ars-grin.gov/gringlobal/query/summary (2023).
Sawler, J. et al. The genetic structure of marijuana and hemp. PLoS One 10, e0133292 (2015).
Lynch, R. C. et al. Genomic and chemical diversity in Cannabis. CRC Crit. Rev. Plant Sci. 35, 349–363 (2016).
Soorni, A., Fatahi, R., Haak, D. C., Salami, S. A. & Bombarely, A. Assessment of genetic diversity and population structure in Iranian Cannabis germplasm. Sci. Rep. 7, 15668 (2017).
Ren, G. et al. Large-scale whole-genome resequencing unravels the domestication history of Cannabis sativa. Sci. Adv. 7 (2021).
Chen, X. et al. Whole-genome resequencing of wild and cultivated cannabis reveals the genetic structure and adaptive selection of important traits. BMC Plant Biol. 22, 371 (2022).
Alsaleh, A. & Yılmaz, G. Exploring Cannabidiol variations, investigation of genetic diversity, population structure and unveiling male-specific genetic marker in industrial hemp (Cannabis sativa L). Genet. Resour. Crop Evol. https://doi.org/10.1007/s10722-024-02015-1 (2024).
Halpin-McCormick, A. et al. Examining population structure across multiple collections of Cannabis. Genet. Resour. Crop Evol. https://doi.org/10.1007/s10722-024-01928-1 (2024).
Carlson, C. H. et al. Morphometric relationships and their contribution to biomass and cannabinoid yield in hybrids of hemp (Cannabis sativa). J. Exp. Bot. 72, 7694–7709 (2021).
Busta, L. et al. Chemical and genetic variation in feral Cannabis sativa populations across the Nebraska climate gradient. Phytochemistry 200, 113206 (2022).
Woods, P., Price, N., Matthews, P. & McKay, J. K. Genome-wide polymorphism and genic selection in feral and domesticated lineages of Cannabis sativa. G3 Genes Genomes Genet. 13 (2023).
Wenger, J. P. et al. Validating a predictive model of cannabinoid inheritance with feral, clinical, and industrial Cannabis sativa. Am. J. Bot. 107, 1423–1432 (2020).
Greenbaum, G., Templeton, A. R., Zarmi, Y. & Bar-David, S. Allelic richness following population founding Events—A stochastic modeling framework incorporating gene flow and genetic drift. PLoS One 9, e115203 (2014).
Miller, R. L. Hemp as a Crop for Missouri Farmers: Markets, Economics, Cultivation, Law. Report to Agriculture Task Force, Missouri House of Representatives (1991).
Cherney, J. & Small, E. Industrial hemp in North america: Production, politics and potential. Agronomy 6, 58 (2016).
Humphrey, J. R. Marketing HempVol. 221, 26–43 (Kentucky Agricultural Experimental Station, 1919).
Husbands, J. D. Feasibility of industrial hemp production in the united States Pacific Northwest. USDA Bull. 153, 42–42 (1909).
Serrote, C. M. L., Reiniger, L. R. S., Silva, K. B., dos Rabaiolli, S. M., Stefanel, C. M. & S. & Determining the polymorphism information content of a molecular marker. Gene 726, 144175 (2020).
Benkirane, C. et al. Population structure and genetic diversity of Moroccan cannabis (Cannabis sativa L.) germplasm through simple sequence repeat (SSR) analysis. Genet. Resour. Crop Evol. 71, 2037–2051 (2024).
Hennink, S. Optimisation of breeding for agronomic traits in fibre hemp (Cannabis sativa L.) by study of parent-offspring relationships. Euphytica 78, 69–76 (1994).
Roger Adams, M., Clark & J. H. Hunt. Structure of cannabidiol, a product isolated from the Marihuana extract of Minnesota wild hemp. I. J. Am. Chem. Soc. 62, 196–200 (1940).
Mechoulam, R. A. et al. Some aspects of cannabinoid chemistry. In The Botany and Chemistry of Cannabis 95–115 (1970).
Haney, A. & Bazzaz, F. A. Some ecological implications of the distribution of hemp (Cannabis sativa L.) in the united States of America. In The Botany and Chemistry of Cannabis 39–48 (1970).
Toth, J. A. et al. Development and validation of genetic markers for sex and cannabinoid chemotype in Cannabis sativa L. GCB Bioenergy 12, 213–222 (2020).
Vergara, D. et al. Gene copy number is associated with phytochemistry in Cannabis sativa. AoB Plants 11 (2019).
Mabry, M. E. et al. Building a feral future: Open questions in crop ferality. Plants People Planet. 5, 635–649 (2023).
Ford, T., Aina, A., Ellison, S., Gordon, T. & Stansell, Z. Utilizing digitized occurrence records of Midwestern feral Cannabis sativa to develop ecological niche models. Ecol. Evol. 14 (2024).
Elshire, R. J. et al. A robust, simple genotyping-by-sequencing (GBS) approach for high diversity species. PLoS One 6, e19379 (2011).
Glaubitz, J. C. et al. TASSEL-GBS: A high-capacity genotyping by sequencing analysis pipeline. PLoS One 9, e90346 (2014).
Li, H. & Durbin, R. Fast and accurate short read alignment with Burrows–Wheeler transform. Bioinformatics 25, 1754–1760 (2009).
Purcell, S. et al. PLINK: A tool set for whole-genome association and population-based linkage analyses. Am. J. Hum. Genet. 81, 559–575 (2007).
Alexander, D. H., Novembre, J. & Lange, K. Fast model-based estimation of ancestry in unrelated individuals. Genome Res. 19, 1655–1664 (2009).
R Core Team. R Core Team. R: A Language and Environment for Statistical Computing (R Foundation for Statistical Computing, 2023).
Wickham, H. Ggplot2: Elegant Graphics for Data Analysis (Springer, 2016).
Excoffier, L., Smouse, P. E. & Quattro, J. M. Analysis of molecular variance inferred from metric distances among DNA haplotypes: Application to human mitochondrial DNA restriction data. Genetics 131, 479–491 (1992).
de Meeûs, T. & Goudet, J. A step-by-step tutorial to use hierfstat to analyse populations hierarchically structured at multiple levels. Infect. Genet. Evol. 7, 731–735 (2007).
ElSohly, M. A. et al. Potency trends of delta9-THC and other cannabinoids in confiscated marijuana from 1980–1997. J. Forensic Sci. 45, 24–30 (2000).
