Genetic Resources and Crop Evolution

, Volume 67, Issue 3, pp 703–714 | Cite as

A multivariate analysis of morphological divergence of “seeds” (achenes) among ruderal, fibre, oilseed, dioecious/monoecious and marijuana variants of Cannabis sativa L.

  • Steve G. U. Naraine
  • Ernest Small
  • Andrew E. Laursen
  • Lesley G. CampbellEmail author
Research Article


Cannabis sativa has been domesticated for stem fibre and oilseed (the two classes are both low in the euphoric cannabinoid THC and called “hemp”), and marijuana (high in THC), and also occurs as weedy, ruderal plants. Achenes (“seeds”) from herbarium collections representative of these classes were assessed for morphological characters and pericarp resistance to fracture. In contrast to ruderal plants, domesticated plants (both hemp and marijuana) possessed achenes that were significantly longer, heavier, covered with a less adherent perianth, and lacking a pronounced basal attenuation. All of these characteristics reflect traits that are advantageous in domesticated plants and are consistent with the “domestication syndrome” found in propagules of other crops. Marijuana achenes, in comparison with hemp achenes, tended to be about 26% shorter and about 32 shades darker (on a 256-bit grayscale). Achenes of fibre cultivars proved to be about 19% longer than the achenes of oilseed cultivars. Achenes of dioecious oilseed cultivars proved to be about 6% longer than the achenes of monoecious oilseed cultivars. The pericarps of hemp seeds were about 26% and about 15% more resistant to fracture than those of ruderal and marijuana plants, respectively.


Achenes Domestication Cannabis sativa Ditchweed Hemp Marijuana 



We thank Brenda Brookes for assistance with figures; Ryerson University Department of Physics, for force measuring software and equipment; Mike Neiser, for building the seed force press; and Michelle Dupuis, for sharing observations.


The authors gratefully acknowledge the funding support from the Natural Sciences and Engineering Research Council (NSERC) Discovery (no. 402305-2011 to LGC) as well as the personal funds of SGU Naraine to build the force meter.

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest on the content of manuscript and study undertaken.


  1. Amaducci S, Coluzzi M, Zatta A, Venturi G (2008) Flowering dynamics in monoecious and dioecious hemp genotype. J Ind Hemp 13:5–19Google Scholar
  2. Anderson EA, Williams L (1954) Maize and sorghum as a mixed crop in Honduras. Ann Mo Bot Gard 41:213–221Google Scholar
  3. Clarke R, Merlin M (2013) Cannabis: evolution and ethnobotany. University of California Press, BerkleyGoogle Scholar
  4. Dufresnes C, Jan C, Bienert F et al (2017) Broad-scale genetic diversity of Cannabis for forensic applications. PLoS ONE 12:e0170522PubMedPubMedCentralGoogle Scholar
  5. Fuller DQ, Allaby R (2009) Seed dispersal and crop domestication: shattering, germination and seasonality in evolution under cultivation. Annu Plant Rev 38:238–295Google Scholar
  6. Funatsuki H, Suzuki M, Hirose A et al (2014) Molecular basis of a shattering resistance boosting global dissemination of soybean. Proc Natl Acad Sci 111:17797–17802PubMedGoogle Scholar
  7. Gilmore S, Peakall R, Robertson J (2003) Short tandem repeat (STR) DNA markers are hypervariable and informative in Cannabis sativa: implications for forensic investigations. Forensic Sci Int 131:65–74PubMedGoogle Scholar
  8. Green AG, Marshall DR (1981) Variation for oil quantity and quality in linseed (Linum usitatissimum). Aust J Agric Res 32:599–607Google Scholar
  9. Gressel J (2005) Crop ferality and volunteerism. CRC Press, Boca RatonGoogle Scholar
  10. Hammer K (1984) The domestication syndrome (in German). Kulturpflanze 32:11–34Google Scholar
  11. Harlan JR (1992) Crops and man. ASA, MadisonGoogle Scholar
  12. Harlan JR (1995) Agricultural origins and crop domestication in the mediterranean region. Diversity 2:14–16Google Scholar
  13. Heredia SM, Ellstrand NC (2014) Novel seed protection in the recently evolved invasive, California wild radish, a hybrid Raphanus sp. (Brassicaceae). Am J Bot 101:2043–2051PubMedGoogle Scholar
  14. Janischevsky DE (1924) A form of hemp in wild areas of Southeastern Russia. Učenye zapiski Saratovskogo Gosudarstvennogo imeni N.G. Černyševskogo Universiteta 2:3–17Google Scholar
  15. Jiang HE, Li X, Zhao YX et al (2006) A new insight into Cannabis sativa (Cannabaceae) utilization from 2500-year-old Yanghai Tombs, Xinjiang, China. J Ethnopharmacol 108:414–422PubMedGoogle Scholar
  16. Kluyver TA, Charles M, Jones G et al (2013) Did greater burial depth increase the seed size of domesticated legumes? J Exp Bot 64:4101–4108PubMedGoogle Scholar
  17. Konishi S, Izawa T, Lin SY et al (2006) An SNP caused loss of seed shattering during rice domestication. Science 312:1392–1396PubMedGoogle Scholar
  18. Li L-F, Olsen KM (2016) To have and to hold: selection for seed and fruit retention during crop domestication. Curr Top Dev Biol 119:63–109PubMedGoogle Scholar
  19. McPartland JM, Naraine SG (2018) Experimental endozoochory of Cannabis sativa achenes. Med Cannabis Cannabinoids 1:96–103. CrossRefGoogle Scholar
  20. McPartland JM, Guy GW, Hegman W (2018) Cannabis is indigenous to Europe and cultivation began during the Copper or Bronze age: a probabilistic synthesis of fossil pollen studies. Veg Hist Archaeobot 2018:1–14Google Scholar
  21. Meyer SE, Carlson SL (2001) Achene mass variation in Ericameria nauseosus (Asteraceae) in relation to dispersal ability and seedling fitness. Funct Ecol 15:274–281Google Scholar
  22. Meyer RS, DuVal AE, Jensen HR (2012) Patterns and processes in crop domestication: an historical review and quantitative analysis of 203 global food crops. New Phytol 196:29–48PubMedGoogle Scholar
  23. Ogg AG, Parker R (1989) Control of volunteer crop plants. Washington State University Extension Bull. EB 1523Google Scholar
  24. Pianka ER (1970) On R- and K- Selection. Am Nat 104:592–597Google Scholar
  25. Porter SS (2013) Adaptive divergence in seed color camouflage in contrasting soil environments. New Phytol 197:1311–1320PubMedGoogle Scholar
  26. Purugganan MD, Fuller DQ (2009) The nature of selection during plant domestication. Nature 457:843–848PubMedGoogle Scholar
  27. Sakuma S, Salomon B, Komatsuda T (2011) The domestication syndrome genes responsible for the major changes in plant form in the Triticeae crops. Plant Cell Physiol 52:738–749PubMedPubMedCentralGoogle Scholar
  28. Serebriakova TY, Sizov IA (1940) Cannabinaceae Lindl. In: Vavilov NI (ed) Kilturnaja Flora SSSR (Flora of Cultivated Plants), vol 5. State Printing Office, Moscow-Leningrad, Kolos, pp 1–53Google Scholar
  29. Small E (1972) Interfertility and chromosomal uniformity in Cannabis. Can J Bot 50:1947–1949Google Scholar
  30. Small E (1974) Morphological variation of achenes of Cannabis. Can J Bot 53:978–987Google Scholar
  31. Small E (1978) A numerical taxonomic analysis of the Daucus carota complex. Can J Bot 56:248–276Google Scholar
  32. Small E (1984) Hybridization in the domesticated-weed-wild complex. In: Grant WF (ed) Plant biosystematics. Academic Press, Toronto, pp 195–210Google Scholar
  33. Small E (2015) Evolution and classification of Cannabis sativa (marijuana, hemp) in relation to human utilization. Bot Rev 81:189–294Google Scholar
  34. Small E (2016) Cannabis: a complete guide. Tayor & Francis/CRC Press, Boca RatonGoogle Scholar
  35. Small E (2017) Classification of Cannabis sativa in relation to agricultural, biotechnological, medical and recreational utilization. In: Chandra S, Lata H, ElSohly MA (eds) Cannabis sativa L.: botany and biotechnology. Springer, Berlin, pp 1–62Google Scholar
  36. Small E (2018) Dwarf germplasm: the key to giant Cannabis hempseed and cannabinoid crops. Genet Resour Crop Evol 65:1071–1107Google Scholar
  37. Small E, Antle T (2003) A preliminary study of pollen dispersal in Cannabis sativa in relation to wind direction. J Ind Hemp 8:37–50Google Scholar
  38. Small E, Beckstead HD (1973) Common cannabinoid phenotypes in 350 stocks of Cannabis. Lloydia 36:144–165PubMedGoogle Scholar
  39. Small E, Cronquist A (1976) A practical and natural taxonomy for Cannabis. Taxonomy 25:405–435Google Scholar
  40. Small E, Marcus D (2003) Tetrahydrocannabinol levels in hemp (Cannabis sativa) germplasm resources. Econ Bot 57:545–558Google Scholar
  41. Small E, Pocock T, Cavers PB (2003) The biology of Canadian weeds. 119. Cannabis sativa L. Can J Plant Sci 83:217–237Google Scholar
  42. Stephens SG (1965) The effects of domestication on certain seed and fiber properties of perennial forms of cotton, Gossypium hirsutum L. Am Nat 99:355–372Google Scholar
  43. Stevens M (2007) Predator perception and the interrelation between different forms of protective coloration. Proc R Soc B Biol Sci 274:1457–1464Google Scholar
  44. Terral J-F, Tabard E, Bouby L et al (2010) Evolution and history of grapevine (Vitis vinifera) under domestication: new morphometric perspectives to understand seed domestication syndrome and reveal origins of ancient European cultivars. Ann Bot 105:443–455. CrossRefPubMedGoogle Scholar
  45. van der Meij MAA, Bout RG (2004) Scaling of jaw muscle size and maximal bite force in finches. J Exp Biol 207:2745–2753PubMedGoogle Scholar
  46. Vavilov NI (1922) Пoлeвыe кyльтypы Югo-Bocтoкa (Field crops of the southeast). Tpyды пo пpиклaднoй бoтaникe, гeнeтикe и ceлeкции (Bull Appl Bot Genet Plant Breed) 13:147–148Google Scholar
  47. Vavilov NI (1931) The role of Central Asia in the origin of cultivated plants. Bull Appl Bot Genet Plant Breed 26:3–44Google Scholar
  48. Venable DL (1992) Size-number trade-offs and the variation of seed size with plant resource status. Am Nat 140:287–304Google Scholar

Copyright information

© Springer Nature B.V. 2019

Authors and Affiliations

  1. 1.Department of Chemistry and BiologyRyerson UniversityTorontoCanada
  2. 2.Ottawa Research and Development Centre, Agriculture and Agri-Food CanadaCentral Experimental FarmOttawaCanada

Personalised recommendations