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The formal botanical taxonomy of Cannabis sativa Linnaeus and C. indica Lamarck has become entangled and subsumed by a new vernacular taxonomy of “Sativa” and “Indica.” The original protologues (descriptions, synonymies, and herbarium specimens) by Linnaeus and Lamarck are reviewed. The roots of the vernacular taxonomy are traced back to Vavilov and Schultes, who departed from the original concepts of Linnaeus and Lamarck. The modified concepts by Vavilov and Schultes were further remodeled by underground Cannabis breeders in the 1980s and 1990s. “Sativa” refers to plants of Indian heritage, in addition to their descendants carried in a diaspora to Southeast Asia, South- and East Africa, and even the Americas. “Indica” refers to Afghani landraces, together with their descendants in parts of Pakistan (the northwest, bordering Afghanistan). Phytochemical and genetic research supports the separation of “Sativa” and “Indica.” But their nomenclature does not align with formal botanical C. sativa and C. indica based on the protologues of Linnaeus and Lamarck. Furthermore, distinguishing between “Sativa” and “Indica” has become nearly impossible because of extensive cross-breeding in the past 40 years. Traditional landraces of “Sativa” and “Indica” are becoming extinct through introgressive hybridization. Solutions for reconciling the formal and vernacular taxonomies are proposed.
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Chapter 4
Cannabis sativa and Cannabis indica
versus Sativaand Indica
John M. McPartland
Abstract The formal botanical taxonomy of Cannabis sativa Linnaeus and C.
indica Lamarck has become entangled and subsumed by a new vernacular taxon-
omy of Sativaand Indica.The original protologues (descriptions, synonymies,
and herbarium specimens) by Linnaeus and Lamarck are reviewed. The roots of the
vernacular taxonomy are traced back to Vavilov and Schultes, who departed from
the original concepts of Linnaeus and Lamarck. The modied concepts by Vavilov
and Schultes were further remodeled by underground Cannabis breeders in the
1980s and 1990s. Sativarefers to plants of Indian heritage, in addition to their
descendants carried in a diaspora to Southeast Asia, South- and East Africa, and
even the Americas. Indicarefers to Afghani landraces, together with their
descendants in parts of Pakistan (the northwest, bordering Afghanistan).
Phytochemical and genetic research supports the separation of Sativaand
Indica.But their nomenclature does not align with formal botanical C. sativa and
C. indica based on the protologues of Linnaeus and Lamarck. Furthermore, dis-
tinguishing between Sativaand Indicahas become nearly impossible because
of extensive cross-breeding in the past 40 years. Traditional landraces of Sativa
and Indicaare becoming extinct through introgressive hybridization. Solutions
for reconciling the formal and vernacular taxonomies are proposed.
4.1 Introduction
Taxonomy includes classication (the identication and categorization of organ-
isms) and nomenclature (the naming and describing of organisms). The formal
botanical taxonomy of Cannabis by Small and Cronquist (1976) recognizes two
subspecies: C. sativa subsp. sativa, and C. sativa subsp. indica. They are considered
different species, C. sativa and C. indica, by some botanists (e.g., Hillig and
Mahlberg 2004; Clarke and Merlin 2013).
J.M. McPartland (&)
GW Pharmaceuticals Place, 1 Cavendish Place, London W1G 0QF, UK
©Springer International Publishing AG 2017
S. Chandra et al. (eds.), Cannabis sativa L. - Botany and Biotechnology,
DOI 10.1007/978-3-319-54564-6_4
In the worlds of recreational and medicinal cannabis, everyone seems to be
talking about Sativaand Indica.This vernacular taxonomy of drug-type
Cannabis has gone viral. Enter Sativa versus Indicainto Google, and the search
returns 45,000 hits. Please stay alert to the fact that Sativaand Indicain
quotation marks are not the same as C. sativa and C. indica written in italics.
Sativaand Indicahave become sources of confusion (Small 2007; Erkelens and
Hazekamp 2014; McPartland 2014; Russo 2016). Hazekamp and Fischedick (2012)
call for an alternative approach, from cultivar to chemovar,where plants are
identied by their chemical ngerprint, rather than a whimsical name.
The goals for this chapter are four-fold: (1) review the formal botanical taxon-
omy of C. sativa and C. indica; (2) trace the history of vernacular Sativaand
Indicaand their misalignment with C. sativa and C. indica; (3) recognize dif-
ferences between Sativaand Indicain phytochemistry and genetics; (4) align
the vernacular taxonomy with the formal botanical taxonomy.
4.2 Formal Botanical Nomenclature: C. sativa
Linnaeus named C. sativa in Species Plantarum, the starting point for botanical
nomenclature (Linnaeus 1753). C. sativa in the strict sense, sensu stricto, is
demarcated by Linnaeuss protologue. The International Code of Nomenclature
(ICN) denes a protologue as everything associated with a taxonomic name at its
rst valid publication. It includes the speciess description, synonymy, and
herbarium specimens (McNeill 2012).
Linnaeuss protologue of C. sativa is described in full for the rst time by
McPartland and Guy (2017). It is abstracted here: Linnaeuss description was
exceptionally brief: a generic account of ower parts, which applies equally to any
plant ever describe in the genus Cannabis (Linnaeus 1753,1754). Linnaeus listed
four synonyms: C. foliis digitatis, C. mas, C. erratica, C. femina; and ve authors
who used those names: himself, Dalibard, van Royen, dAléchamps, and Bauhin.
The authors and their synonyms delimit C. sativa to plants from northern Europe.
His herbarium specimens also came from northern Europe. Linnaeuss type
specimen of C. sativa is stored at the Linnaeus herbarium (Fig. 4.1). The seeded
pistillate plants morphology is consistent with a northern European ber-type
landrace. Its inorescences are loose, not dense; subtending oral leaves have a
sparse covering of sessile glandular trichomes; perigonal bracts that enclose ach-
enes (seeds) have a relatively sparse covering of capitate stalked glandular
(CSG) trichomes. Evidence by Stern (1974) indicates that Linnaeus collected the
specimen in Sweden. Other C. sativa specimens collected by Linnaeus and stored at
the British Museum are consistent with the old cultivated hemp stock of northern
Europe(Stern 1974).
Linnaeus notably excluded Asian plants from the C. sativa protologue. He
certainly knew about Asian Cannabis. Sixteen years earlier, Linnaeus (1737) cited
six authors who assigned names to psychoactive Asian Cannabis: C. Bauhin
102 J.M. McPartland
(Cannabi similis exotica), J. Bauhin and Cherler (Bangue cannabi simile), Ray
(Bangue cannabi), Rheede (Kalengi cansjava and Tsjeru cansjava), Morison
(Cannabis peregrina gemmis fructuum longioribus), and Kaempfer (Ba and Ma). In
summary, LinnaeussC. sativa taxon represents rope, not dope(McPartland et al.
2000). It does not align with vernacular Sativa,known for its potent
4.3 Formal Botanical Nomenclature: C. indica
Lamarck (1785) coined C. indica for plants of Indian provenance and their
descendants in Southeast Asia and South Africa. For a full account of his proto-
logue see McPartland and Guy (2017). The description of C. indica differed from
that of C. sativa by eight very distinctmorphological characters, in stalks,
branching habitus, leaets, and owers. Lamarck noted ne details in C. indica,
female owers have a vellous calyx and long styles.In other words, the perigonal
bract (calyx) is velvety (vellous), due to a dense pubescence of CSG trichomes.
Nearly 230 years passed before others noted long styles in C. indica (Small and
Naraine 2015a). Lamarck also described chemotaxonomic differences: C. indica
produced a strong odor, and caused intoxication when smoked in a pipe.
Lamarcks type specimen at the Paris herbarium was collected by Pierre
Sonnerat, probably around Pondicherry. Lamarcks specimen shows denser growth
and more compact branching than Linnaeuss specimen (Fig. 4.1). Its inorescences
Fig. 4.1 Herbarium type specimens of C. sativa L. (left), and C. indica Lam. (right), photographs
courtesy of McPartland and Guy (2017)
4Cannabis sativa and Cannabis indica 103
are somewhat dense; subtending oral leaves have an abundant covering of sessile
glandular trichomes; perigonal bracts express a moderate density of CSG trichomes.
The styles and stigmas are prominent, agglutinized, and light brown in color.
Lamarck listed six synonyms: Cannabi similis exotica Bauhin (who cited da
Orta and Acosta in Goa, India); Kalengi-cansjava and Tsjeru-cansjava Rheede
(plants from Kochi, India); Cannabis peregrina gemmis fructuum longioribus
Morison (who cited Bauhin and Rheede); C. indica Rumph (plants from Indonesia);
and Dakka ou Bangua Prévost (who cited Kolb in South Africa). In summary,
Lamarck delimited C. indica to plants from southern India and their descendants in
Indonesia and South Africa.
4.4 The Slide from Formal to Vernacular
How did the species names C. sativa and C. indica reappear inaccurately as
Sativaand Indica? We traced a path through Afghanistan by Nikolai Ivanovich
Vavilov and Richard Evans Schultes. Vavilov traveled there in 1924, where he
encountered Afghani farmers who cultivated Cannabis for gashisha (hashīsh). He
assigned these plants to C. sativa (Vavilov and Bukinich 1929). This departed from
Linnaeuss concept of C. sativa as a European ber-type plant. Vavilov also
encountered wild-type and feral plants, which he named, respectively, C. indica
var. karistanica and C. indica f. afghanica. His student Tatiana Yakovlevna
Serebriakova assigned Afghani plants to C. sativa, and Indian plants to C. indica
(Serebriakova and Sizov 1940).
Schultes travelled to Afghanistan in 1971. Schultes et al. (1974) narrowly typ-
ied C. indica to plants in Afghanistan, with broad, oblanceolate leaets, densely
branched, with very dense inorescences, more or less conical in shape, and very
short (<1.3 m). This departed from the original taxonomic concept of Lamarck,
who was entirely unfamiliar with Afghani Cannabis. Lamarcksindica designates
Cannabis from Indiarelatively tall, laxly branched, with narrow leaets.
Anderson (1980) echoed Schultes and assigned Afghani plants to C. indica
short, conical, densely branched, with broad leaets. He assigned plants from India
to C. sativarelatively tall, laxly branched, with narrow leaetsplants that
Lamarck would have called C. indica. Anderson illustrated these concepts in a line
drawing (Fig. 4.2) that now appears everywhere on the internet.
Clarke (1981) referred to plants from Afghanistan as type examples for
Cannabis indica.Cherniak (1982) assigned cannabis sativato plants of South
Asian heritage (Nepal, Burma, Thailand), and their descendants in Columbia,
Jamaica, and Mexico. He applied the name cannabis indicato plants from
Afghanistan. His classication gets a bit muddled, because he also categorizes
plants from India as cannabis indica.The earliest consistent use of Sativaand
Indicaappears in a Dutch seed catalog (Watson 1985).
104 J.M. McPartland
Meijer and van Soest (1992) brought attention to this vernacular taxonomy in
peer-reviewed literature: Indicarefers to plants with broad leaets, compact habit,
and early maturation, typied by plants from Afghanistan. Sativarefers to plants
with narrow leaets, slender and tall habit, and late maturation, typied by plants
from India and their descendants in Thailand, South and East Africa, Colombia, and
Clinical descriptions of Sativaand Indicaare barely a decade old (Corral
2001; Black and Capler 2003): Sativaplants produce much more
hydrocannabinol (THC) than cannabidiol (CBD), and produce a terpenoid prole
that smells herbalor sweet.”“Sativaimparts a stimulating, uplifting, and
energizing psychoactivity, and is recommended for treating depression, headaches,
nausea, and loss of appetite. Indicaplants produce a nearly equal THC-to-CBD
ratio, and a terpenoid prole that imparts an acrid or skunkyaroma. Indica
induces relaxing, sedating, and pain-reducing effects, and is suggested for treating
insomnia, pain, inammation, muscle spasms, epilepsy, and glaucoma.
McPartland et al. (2000) separated Sativaand Indicafrom European hemp,
and provisionally named the three populations C. indica, C. afghanica, and C.
sativa, respectively. Small (2007) noted that Sativaand Indicawere quite
inconsistentwith formal nomenclature, because C. sativa subsp. sativa applied to
non-intoxicant plants.
Hillig (2004a,b,2005a,b) avoided formal/vernacular conicts by applying the
name narrow-leaet drug (NLD) biotypeto plants corresponding with Lamarcks
C. indica. He assigned wide-leaet drug (WLD) biotypeto plants corresponding
with Vavilovsafghanica (i.e., SchultessC. indica and vernacular Indica). This
nomenclature has gained traction (e.g., McPartland and Guy 2004; Russo 2007;
Lynch et al. 2015).
Fig. 4.2 Cannabis
vernacular taxonomy, image
adapted from Anderson
4Cannabis sativa and Cannabis indica 105
McPartland and Guy (2004) and McPartland (2014) proposed reconciling
Sativaand Indicawith C. sativa and C. indica by correcting the vernacular
nomenclature: Sativais really indica, and Indicais actually afghanica, and
Ruderalisis usually sativa. The initial reaction to this proposition by recreational
users was negative. An editorial in High Times characterized the corrected
nomenclature as undoubtedly a little kooky(Sirius 2015).
Researchers, however, are starting to take it on board (e.g, Henry 2015). Clarke
and Merlin (2016) published a vernacular correction nearly identical to McPartland
(2014), although they did not cite the precedent publication. Two table headings in
their respective taxonomic tables are exampled:
McPartland (2014): Indica (formerly Sativa)
Clarke and Merlin (2016): IndicaWrongly called sativa.
The title of their article is adapted from other antecedents, also uncited
(Tejkalováand Hazekamp 2014; Piomelli and Russo 2016). If these phrases were
botanical names, a taxonomist would invoke the principle of priority. For example,
Clarke and Merlin (2013) erected a new biotype name, BLD(broad leaf drug).
They objected to Hilligs names of biotypes based on leaf shape. Nevertheless,
invoking priority, BLD is a later synonym of Hilligs WLD. Similarly, Clarke and
Merlin (2015) strenuously rejected Smalls taxonomic character intoxicant.They
replaced it with psychoactive.
Erkelens and Hazekamp (2014) outlined the history of Indicaand they
emphasized taxonomical conicts between monotypic and polytypic views of
Cannabis. For the rest of this chapter, Sativarefers to the NLD biotype, or plants
of Indian heritage (including their putative descendants in Southeast Asia, Africa,
and the Americas). Indicarefers to the WLD biotype, or Afghani landraces
(including related populations in northwestern Pakistan bordering Afghanistan, and
possibly neighboring TurkestanUzbekistan and Xīnjiāng).
4.5 The Hybridization Impasse
Selective cross-breeding of drug-type Cannabis accelerated in the 1970s.
Germplasm from Afghanistan was smuggled into California in the early 1970s (D.
Watson, pers. commun. 1984), or the late 1970s (Clarke 1987). During the 1980s at
least seven Cannabis breeders sold exotic germplasm in Holland. They crossed
plants of Indian heritage (sweet but late maturing) with Afghani landraces, valued
for rapid maturation, cold-tolerance, short stature, and dense, tightly-packed ower
clusters. By the late 1980s, nearly all drug-type Cannabis grown in the USA,
Canada, and Europe had been hybridized. Unadulterated plants of Indian heritage
and Afghani landraces became difcult to obtain (Clarke 1987).
Alarmingly, foreign germplasm has corrupted Indian and Afghani landraces in
their former centers of diversity. Peterson (2009) deplored the importation of
106 J.M. McPartland
Skunk #1into South Africa around 1984, which destabilizedthe genepool.
Jamaicans have replaced gañjāof Indian origin with Afghani hybrids (J. McP, pers.
observ. 2013). Beisler (2006) boasted of importing and growing Mexican Goldin
Afghanistan around 1972. Pietri (2009) stated that Beisler crossed Acapulco Gold
with Afghani landraces. Turner et al. (1979b) analyzed 12 accessions collected in
northwest India, and some plants in Punjab expressed low THC/CBD proles
suggestive of Afghani landraces.
Ubiquitous hybridization of Sativaand Indicahas rendered their distinctions
almost meaningless. Most hybrids are characterized as Sativa-dominantor
Indica-dominant.The arbitrariness of these designations is illustrated by
AK-47,a hybrid that won Best Sativain the 1999 Cannabis Cup, and won
Best Indicafour years later. Hybrids have been assigned strainnames. The
desire for unique weed has led to an explosion of new strain names. At the dawn of
this era, Watson (1985) offered 10 strains for sale. Fifteen years later, Clarke (2001)
estimated that Dutch seed companies offered 150 strains for sale, and 80% of them
contained hybridized ancestry from Watsons original strains. A decade later the
number of named strains reached 900 (Cannabis Strain Database 2010). Leay
(2015) listed 1535 strain names, and Seednder (2015) listed 6510 strain names.
Doyle (2007) called the strain names ganjanyms.
In todays largely illicit market, strain names are swapped and counterfeited, and
generally unreliable (Lee 2013; Sawler et al. 2015; Pierson 2016). Unrecognized
hybrids have plagued recent taxonomic studies of Sativaand Indica.
Unrecognized hybrids assigned to C. sativa or C. indica dampen signal in any
taxonomic methodology. Widespread crossbreeding and introgression make it
challenging to meet the third goal of this book chapter: identifying differences
between the NLD biotype (Sativa) and the WLD biotype (Indica). The biotypes
show differences in cannabinoids, terpenoids, and genetics.
In the next couple sections of this chapter, analytical studies that measured
cannabinoids and terpenoids in NLD and WLD biotypes will be compared. This
comparison is hampered by the fact that different studies used different analytical
methods (e.g., gas chromatography versus high performance liquid chromatogra-
phy). These analytical methods may vary in their yields of cannabinoids and ter-
penoids (Wheals and Smith 1975; Hazekamp et al. 2005; Giese et al. 2015).
Cannabinoid and terpenoid content is best measured in a common garden experi-
ment (CGE), where plants from different places are grown in a single location,
under identical environmental conditions, and uniformly processed.
4.6 Cannabinoids
Cannabinoid content differs in terms of quantity and quality; these differ in their
modes of inheritance (Hillig 2002). Cannabinoid quantity (dry weight percentage)
is polygenic and inuenced by environmental factors. Cannabinoid quality (the
THC/CBD ratio, known as the cannabinoid prole or chemotype) is largely genetic
4Cannabis sativa and Cannabis indica 107
possibly monogenic. For more information regarding the differences between
quantity and quality, see this books chapter by Grassi and McPartland. The
THC/CBD ratio, a dimentionless ratio, cancels two quantities (THC%, CBD%), and
provides a more valid comparison between plants cultivated in different
Hillig and Mahlberg (2004) published an exemplary CGE: 157 Cannabis
accessions from around the world, with passport data regarding provenance.
Accessions were classied into seven biotypes, with a priori segregation based on
geographic origins and a genetic analysis (Hillig 2005a). We will focus on two
populations in their study: the WLD biotype (n = 12 from Afghanistan, the
North-West Frontier Province of Pakistan, and Uzbekistan) and the NLD biotype
(n = 27 from India, Thailand, Cambodia, Mexico, Colombia, Jamaica, South- and
East Africa).
They prepared a voucher specimen of each accession, deposited in a herbarium.
Voucher specimens are critical for authenticating the identication of a taxon;
vouchers allow other researchers to retrospectively analyse accessions (Culley
2013). Hillig and Mahlberg made a great effort to exclude hybrids. For example,
Not everything from Afghanistan is Afghani(Hillig, pers. commun., 2006). An
examination of their vouchers reveals that a few hybrids snuck into their analysis.
They cultivated plants in a glasshouse under natural and supplemental light, and
staggered the harvest to sample each accession at peak, uniform maturity.
Cannabinoids were measured in individual plants, rather than in bulked samples.
Results obtained from NLD and WLD biotypes are presented in Table 4.1.
Hillig and Mahlberg reported a statistical difference in CBD%, but no statistical
difference in THC% (Table 4.1). They depicted cannabinoid proles in graphs
(histograms and Cartesian graphs), but they did not present actual numerical data.
We calculated cannabinoid proles of NLD and WLD biotypes from their data in
Table 4.1, as the quotient of THC/CBD. NLD biotype = 5.48/0.02 = 274.0, WLD
biotype = 6.49/1.21 = 5.4. The order of magnitude difference between 274.0 and
5.4 is signicant, although statistical inferences cannot be calculated for n = 2.
The WLD biotype produced a much greater concentration of cannabinoids
(represented by THC% + CBD% in Table 4.1). This is likely due to the WLD
biotypes greater density of perigonal bracts, and greater expression of CSG tri-
chomes on oral leaves, compared to NLD plants. The size of resin heads (gland
heads) may also differ. Small and Naraine (2015b) measured resin head size in ten
strains of high-THC medical marijuana(WLD-NLD hybrids), which averaged
129 µm in diameter. Seven cultivars of low-THC industrial hemp averaged 81 µm.
Table 4.1 Cannabinoid content (mean ±standard deviation) in two Cannabis biotypes, data
from Hillig and Mahlberg (2004)
NLD biotype 5.48 ±2.41 0.02 ±0.02 5.50 ±2.42 0.25 ±0.40
WLD biotype 6.49 ±4.09 1.21 ±2.78 7.70 ±3.45 0.14 ±0.30
Means in this column are statistically different using Students pairwise ttest (p!0.05)
108 J.M. McPartland
Previous studies of CSG trichome density did not include Afghani plants (Small
et al. 1976; Turner et al. 1977).
Hillig and Mahlberg measured cannabigerol (CBG), cannabigerol-
monomethylether (CBGM), and cannabichromene (CBC), with no statistical dif-
ferences between NLD and WLD biotypes. NLD biotypes produced more
tetrahydrocannabivarin (THCV) and cannabidivarin (CBDV) than WLD biotypes.
THCV and CBDV are the short-tailed C
analogs of THC and CBD, respectively.
This trend can be seen in data reported by Turner et al. (1973). Some researchers
include C
analogs in the calculation of cannabinoid proles, as THC
+THCV/CBD+CBDV (Turner et al. 1979a; Onofri et al. 2015; Welling et al. 2016).
Prior to Hillig and Mahlberg, few CGEs studied Afghani landraces. Holley et al.
(1975) analyzed a worldwide collection of females, males, mixtures of females and
males, immature plants, and cross-pollinated hybrids. Here are some of their results,
in nonhybridized females, as THC%, CBD%, and the quotient of THC/CBD:
Afghanistan: 0.59/1.26 = 0.47, India A: 1.78/0.03 = 59.3, India E:
3.31/0.02 = 165.5, Nepal: 2.75/0.02 = 137.5, Pakistan: 1.32/0.01 = 132, South
Africa D: 1.84/0.01 = 184, South Africa E: 0.62/0.06 = 10.3, South Africa F:
0.33/0.01 = 33, and Brazil: 2.16/0.01 = 216. In summary, the results by Holley and
colleagues are similar to those of Hillig and Mahlberg: the THC/CBD ratio in
Afghani landraces was much lower than that of NDL landraces.
Meijer et al. (1992) analyzed 97 accessions, many of uncertain provenance (e.g.,
Nederwiet), or hybridized material, such as Skunk #1.They included three
accessions from Afghanistan: Rjaf 1: 1.15/1.60 = 0.720; Afghanistan:
1.69/0.25 = 6.76; and Afgaan: 2.00/1.18 = 1.69. No samples from India were
included in the study. Two early CGEs that lacked Afghani plants were never-
theless instructive, because they analyzed plants of Indian heritage prior to the era
of widespread hybridization. Fetterman et al. (1971) measured cannabinoid ratio,
including cannabinol (CBN), as THC+CBN/CBD, in samples from Mexico68: 1.0
+0.55/0.075 = 20.5, Mexico69: 1.4+0.073/0.12 = 12.3, Thailand A:
2.2 + trace/0.16 = 13.8, Thailand B: 1.3 + trace/0.11 = 11.8.
Small and Beckstead (1973) analyzed 350 accessions from around the world.
Many accessions came from botanical gardens, of questionable provenance (e.g.,
three indica accessions with no measurable THC). Here are some of their results
accessions of Indian heritage with solid passport data, presented as THC%, CBD%,
and THC/CBD. India: 1.58/0.15 = 10.5, Malawi A: 1.44/0.5 = 28.8, Malawi B:
1.92/0.11 = 17.45, Malawi C: 0.90/0.07 = 12.86, South Africa: 1.34/0.09 = 14.89,
Rhodesia: 0.73/0.06 = 12.17, Cambodia: 1.03/0.12 = 8.5, Uganda: 2.56/0.34 =
7.53, Mauritius: 1.90/0.26 = 7.31, Mexico: 1.52/0.23 = 6.61, Jamaica: 1.19/0.3 =
3.97. No Afghani landraces were included in the study.
Comparisons of police-conscated samples lack the accuracy of CGEs.
However, some studies are instructive because they predate the era of widespread
hybridization. Marshman et al. (1976) tested 36 samples from Jamaica, with a mean
of 3.03/0.10 = 29.5 (two samples reached 99.0 and 104.4). Jenkins and Patterson
(1973) measured THC, CBN, and CBD in herb and hashīsh seizures. Means were
calculated from their raw data: Afghanistan (n = 4): 52.0 + 12.0/36.1 = 1.77,
4Cannabis sativa and Cannabis indica 109
Pakistan (n = 19): 35.7 + 16.1/48.2 = 1.07, South Africa (n = 6): 75.6 +
16.0/8.5 = 10.78, Jamaica (n = 7): 77.5 + 13.4/9.1 = 9.99, Burma (n = 5):
15.7 + 67.9/16.34 = 5.12.
Mobarak et al. (1978) analyzed hashīsh from Kandeh in Petschtal,a.k.a.
Kandai in Pech River valleyjust 55 km from where Vavilov collected C. indica
var. afghanica. They report THC + CBN/CBD as 8 + 14.4/11.6 = 1.93. Martone
et al. (1990) analyzed THC+CBN/CBD in hashīsh seizures, including Afghanistan:
4.45 + 0.36/1.73 = 2.78 and India: 4.48 + 0.40/1.59 = 3.07.
Researchers after Hillig and Mahlberg faced greater difculties parsing hybrids
from their studies. Mahlberg and Hillig collected germplasm during the 1970s
1990s. Since then, unadulterated landraces have become needles in haystacks. For
example, de Meijer and colleagues recently reported Afghani plants with extremely
high THC/CBD quotients (e.g., 683.7 and 516.6, Onofri et al. 2015), or extremely
low THC/CBD quotients (e.g., 0.04, Meijer et al. 2009). These results depart from
an earlier study by Meijer et al. (1992), where THC/CBD quotients for Afghani
plants averaged around 3.1 (Meijer et al. 1992).
Researchers in Holland analyzed 11 strains in a non-CGE study (Fischedick
et al. 2010). No provenance was provided, partially due to proprietary rights. Also,
the operational gray-zone of Dutch coffeeshops (legal front door, illegal back
door) impedes information transfer regarding passport data and provenance. Six
strains were considered nonhybridized Indicas:AD,”“AF,”“AM,”“AN,
AO,and Bedropuur.All six were essentially devoid of CBD. This was a major
departure from studies of Afghani landraces collected in the 1970s1990s, which
had signicant CBD levels. The lack of CBD in 21st century Indicasis incon-
sistent with Afghani landraces from the 1970s1990s.
The same group (Tejkalováand Hazekamp 2014; Tejkalová2015) conducted an
enlarged study of typical representativesof Sativa(n = 44) and Indica
(n = 77). They obtained samples from Dutch coffeeshops and proprietary sources
(Bedrocan BV, HempFlax BV), with limited information regarding provenance.
They used a multivariate clustering method, Principal Component Analysis (PCA).
The PCA scatterplot clearly discriminated Sativasamples from Indicasamples,
but THC and CBD did not provide discriminatory value (i.e., the PCA weights or
eigenvector values for THC and CBD did not discriminate between Sativaand
Hazekamp et al. (2016) adjusted their sample size to Sativa(n = 68) and
Indica(n = 63), obtaining samples from the same sources. This time they pre-
sented a PCA scatterplot as well as numerical means. They found no signicant
differences between Sativaand Indicain either THC or CBD content. Sativa
THC/CBD means 12.74/0.38 = 33.5; IndicaTHC/CBD means 13.71/0.30 =
Elzinga et al. (2015) analyzed 35 strains obtained from chemotypical medicinal
cannabis dispensaries.They assigned strains to Indica,”“Sativa,or Hybrid
based on reports by the Leay website. Instead of THC/CBD ratios, they presented
average THCmax%for each strain. Indica(n = 13) averaged 17.30%, and
Sativa(n = 5) averaged 13.84%. For CBD they offered only summary statistics
110 J.M. McPartland
for all 35 strains: mean 0.6%, median 0.3%. Only one strain produced >1.49%
CBD. Based on such low CBD levels, it can be deduced that all but one of their
strains were high THC hybrids. They say so in a roundabout way, previous papers
used samples collected worldwide, and based upon their reported cannabinoid
levels, are not representative of the cannabi[s] currently available in the United
States to patients and recreational users.Not surprisingly, their PCA analysis does
not support the classication between indica and sativa as it is commonly pre-
sented(i.e., classication by Leay).
Lynch et al. (2015) concatenated databases for a genotype-chemotype study,
which may explain their unique results. They ltered two large databases of strain
sequences, and found 195 strains with common polymorphic sites (see genetics
section). Cluster analysis of polymorphic sites sorted the 195 strains into groups
named WLD and NLD biotypes. The Strain Fingerprintdatabase, developed by
Steep Hill Labs (and displayed by Leay on its website) included chemotype
information for 54 of the 195 strains. Lynch and colleagues presented histograms of
mean THC% and CBD%, from which cannabinoid ratios can be estimated: WLD:
16.5/0.2 = 82.5, NLD: 14.2/2.2 = 6.45. Thus the latest THC/CBD ratios of 21st
century ganjanyms shows a stunning reversal of THC/CBD ratios compared to their
corresponding 1970s1990s landraces.
4.7 Terpenoid Studies
Terpenoids include simple terpenes (isoprenes) and modied terpeneswhere
methyl groups have been moved or removed, or oxygen added as alcohols, esters,
aldehydes, or ketones. The characteristic odor of Cannabis comes from its unique
blend of monoterpenoids (C
templates) and sesquiterpenoids (C
plates). Terpenoids provide a key distinguishing feature between skunkyAfghanis
and herbal-sweetplants of Indian heritage (Black and Capler 2003). Despite this
key diagnostic feature, few terpenoid studies have included Afghani landraces.
Hood and Barry (1978) analyzed headspace”—the odor given off by plants, rather
than contents of glandular trichomes. Headspace favors the detection of monoter-
penoids over less-volatile sesquiterpenoids. Hood and Barry quantied 17 terpenoids in
14 accessions, including plants from Afghanistan and Pakistan (n = 3) and plants from
India and Mexico (n = 5). Running statistics on their raw data revealed some terpenoids
with statistical differences: Hood and Barry reported more limonene in Af/Pak plants
(mean 16.5% ±1.66 SD) than Indi-Mex plants (6.5% ±1.01, p< 0.001), and more
b-farnesene in Indi-Mex (0.44% ±0.13) than Af/Pak (0.10% ±0.05, p= 0.10).
Differences in three other terpenoids fell a little short of statistical signicance: more b-
caryophyllene in Indi/Mex (3.0% ±0.39) than Af/Pak (1.9% ±0.52, p= 0.16), more
a-humulene in Indi/Mex (0.76% ±0.20) than Af/Pak (0.53% ±0.15, p= 0.20), and
more b-myrcene in Af/Pak (10.0% ±0.53) than Indi/Mex (7.6% ±1.3, p= 0.21).
Hillig (2004b) identied 21 terpenoids in a subset of the Cannabis collection that
he analyzed for cannabinoids. He compared terpenoid proles in WLD biotypes
4Cannabis sativa and Cannabis indica 111
(n = 9) and NLD biotypes (n = 21), using a PCA analysis. The PCA scatterplot
clearly discriminated WLD plants from NLD plants. Four terpenoids with the
greatest discriminatory value (i.e., greatest PCA weight or eigenvector value) were
sesquiterpene alcohols: guaiol, c-eudesmol, b-eudesmol, and a peak tentatively
identied as a-eudesmol. All signicant differences (p< 0.05) are presented in
Table 4.2. Regarding b-myrcene, Hillig reported the same trend as Hood and Barry:
WLD = 9.0%, NLD = 5.8%, falling short of statistical signicance.
Fischedick et al. (2010) analyzed 23 terpenoids in six strains considered non-
hybridized Indicas:AD,”“AF,”“AM,”“AN,”“AO,and Bedropuur.As
mentioned above, the Indicastrains had no quantiable CBD, so they likely were
unrecognized hybrids. They also analyzed ve strains classied as hybrids: AE
(mostly Sativa), AG(Indica/Sativa), Ai94(mostly Sativa), Bediol
(Indica/Sativa/Ruderalis), Bedrocan(Indica/Sativa). (Indica dominant).
They made an interesting discovery: three Indicas(i.e, unrecognized hybrids
(Bedropuur,”“AO,and AF) expressed measurable levels of guaiol, c-eudes-
mol, and b-eudesmol. These sesquiterpene alcohols are unique to Afghani landraces
(Hillig 2004b). Furthermore, the same three strains contained higher levels of
limonene than the other accessions, results consistent with Hillig. Thus hybridized
Indicas,despite selection for elevated THC/CBD ratios, retained unique ter-
penoids in common with their landrace ancestors.
Hazekamp and Fischedick (2012) identied terpenoids in more hybrids, two
Sativa dominantsamples (Amnesia,”“Bedrobinal) and two Indica dominant
samples (White Widow,”“Bedica). Once again, only Indica dominanthybrids
contained guaiol, c-eudesmol, and b-eudesmol.
The aforementioned study of samples from Dutch coffeeshops and proprietary
sources (Tejkalováand Hazekamp 2014; Tejkalová2015) analyzed 21 monoter-
penoids and 19 sesquiterpenoids. Multivariate clustering with PCA produced a
scatterplot that segregated Sativaand Indicainto distinct clusters, with some
Table 4.2 Terpenoid
(mean ±standard deviation)
in NDL and WLD biotypes,
reported as statistically
different by Hillig (2004b)
Limonene 1.3% ±1.2 4.0% ±4.3
c-terpinene 0.2% ±0.2 0.1% ±0.2
b-fenchol 0.2% ±0.2 0.8% ±0.9
Terpinoline 4.4% ±8.0 1.0% ±2.9
b-caryophyllene 15.7% ±7.2 9.7% ±6.2
a-guaiene 1.0% ±1.3 0.4% ±0.7
Trans b-farnesene 7.6% ±4.4 4.1% ±3.3
Caryophyllene oxide 8.9% ±7.9 4.2% ±4.2
Guaiol 0.2% ±0.4 3.5% ±1.8
c-eudesmol 0.6% ±0.6 4.8% ±2.1
b-eudesmol 0.8% ±0.6 7.4% ±4.0
a-eudesmol (peak 41) 0.1% ±0.3 1.4% ±1.4
Percentages are ratios of individual peak areas relative to the total
area of all 48 terpenoid peaks
112 J.M. McPartland
overlap and outliers. Indicastrains produced more guaiol, c-eudesmol, and b-
eudesmol, as well as another sesquiterpene alcohola-bisabolol, plus three mon-
terpene alcohols: a-terpineol, b-fenchol, and linalool. Sativastrains leaned
towards unoxygenated sesquiterpenes: a-humulene, trans-b-caryophyllene, a-
guaiene, and trans-a-bergamotene.
Hazekamp et al. (2016) adjusted their sample size to Sativa(n = 68) and
Indica(n = 63), and analyzed 17 monoterpenoids and 19 sesquiterpenoids. Once
again, Indicasamples produced more sesquiterpene alcohols than Sativa
samples (guaiol, c-eudesmol, b-eudesmol, and a-bisabolol), as well as more
monterpene alcohols (a-terpineol, b-fenchol, and linalool). This time they also
report two more monterpene alcohols (cis-sabinene hydrate, borneol) in Indica.
They concluded that hydroxylated terpenoids in general, not just sesquiterpene
alcohols, distinguished Indicastrains. They also report signicantly more limo-
nene and myrcene in Indica,consistent with Hilligs Afghani landraces (although
they erroneously state that Hillig found less myrcene in Afghani landraces).
Mansouri et al. (2011) analyzed terpenoids in Iranian plants, which expressed
signicant amounts of b-eudesmol and c-eudesmol, like plants of Afghani heritage.
Casano et al. (2011) compared 16 unnamed hybrid accessions, characterized as
mostly Indicaor mostly Sativa.”“Mostly Indicaplants produced signicantly
higher levels of limonene, b-myrcene, and camphene. Mostly Sativaproduced
signicantly higher levels of sabinene, -3-carene, a-phellandrene, 1,8-cineole, cis-
b-ocimene, trans-b-ocimene, and a-terpinolene.
Elzinga et al. (2015) assigned strains to Sativaor Indicaaccording to the
Leay database, as described earlier. They noted that strains named Kush,char-
acteristic of the wide leaet drug type strains originating from Hindus Kush region
of Afghanistan and Pakistan,contained higher levels of guaiol, b-eudesmol, b-
myrcene, trans-ocimene, and b-pinene.
Lynch et al. (2015) concatenated databases for a genotype-chemotype study, see
explanation. They reported seven terpenoids in strains assigned to the NLD biotype
(n = 35) or the WLD biotype (n = 17). NLDs produced greater levels of b-myrcene
and a-terpinolene (0.48% and 0.16%, respectively) than did WLDs (0.35% and
0.09%). WLDs produced greater levels of linalool (0.08%) than did NLDs (0.02%).
No statistically signicant differences were seen in limonene, a-pinene, b-car-
yophyllene, and caryophyllene oxide. No sesquiterpene alcohols were measured.
Terpenoids modulate the effects of THC (McPartland and Pruitt 1999;
McPartland and Russo 2001). Two terpenoids in particular have gained attention.
Anonymous (2006) claimed that b-myrcene added to THC made the drug sensation
more physical, mellow, sleepy,whereas limonene added to THC made the drug
sensation more cerebral and euphoric.Russo (2011) attributed the sedative
couch-lockof Indicato b-myrcene, and Russo (2016) attributed the uplifting
effects of Sativato limonene. Chemotype studies do not entirely support these
observations. Regarding limonene, earlier studies showed greater amounts in
Afghani landraces than in plants of Indian heritage (Hood and Barry 1978; Hillig
2004b). This trend was seen in some recent studies of mostly Indica
4Cannabis sativa and Cannabis indica 113
(i.e., Afghani), versus mostly Sativa(i.e., plants of Indian heritage) (Fischedick
et al. 2010; Casano et al. 2011; Hazekamp et al. 2016). Other studies of 21st century
ganjanyms show no differences in limonene between Indicaand Sativa
(Elzinga et al. 2015; Lynch et al. 2015).
Regarding b-myrcene, earlier studies showed greater amounts in Afghani lan-
draces than in plants of Indian heritage, albeit short of statistical signicance (Hood
and Barry 1978; Hillig 2004b). This trend continued in four recent studies of
Indicaversus Sativa(Fischedick et al. 2010; Casano et al. 2011; Elzinga et al.
2015; Hazekamp et al. 2016), although other studies show no differences
(Hazekamp and Fischedick 2012), or even a reversal of earlier results (Lynch et al.
4.8 Genetic Studies
Hillig (2005a) analyzed allozyme variation in the same Cannabis collection tested
for cannabinoids. Samples were evaluated for variation at 17 gene loci, and fre-
quencies of 52 alleles were subjected to PCA. The PCA scatterplot segregated
drug-type plants and ber-type plants into distinct clusters, but the WLD ellipse and
NLD ellipse substantially overlapped.
Gilmore et al. (2007) examined 76 Cannabis accessions for ve polymorphic loci
sequenced from chloroplast and mitochondrial DNA. The studysaws are manifold,
but parsimony analysis recovered three clades. Clade A comprised a majority of
ber-type plants. Clade B included Afghani landraces along with most drug strains
hybrids and police seizures. Clade C was the most interestingnothing but classic
Sativas: 12 landraces from India, Nepal, Thailand, Jamaica, Mexico, and Africa.
Gilmore (2005) gave the name C. sativa rasta to plants in Clade C.
Knight et al. (2010) tested six seized plants, identied by their morphology as
Sativa(n = 2) or Indica(n = 4). Five short tandem repeat (STR) loci, analyzed
with PCA, clearly segregated Sativaplants from three of the Indicaplants. The
fourth Indicaexhibited a unique genotype suggestive of a polyploid condition.
Piluzza et al. (2013) compared 19 accessions: one Afghani, ve of Indian her-
itage, three Skunkhybrids, and an assortment of ber-type plants from Europe
and East Asia. Six RAPD primers detected DNA polymorphisms, with haplotypes
clustered using a neighbor-joining algorithm. Plants of Afghani and Indian heritage
fell into separate clusters. Each shared interesting clade-mates. The Afghani lan-
drace was sister to a cluster of ber-type plants. The cluster of Indian heritage plants
was sister to the Skunkcluster.
Onofri et al. (2015) searched for single nucleotide polymorphisms (SNPs) in
THCA-S sequences. They found nine unique THCA-S sequences amongst 18
accessions of ber- and drug-type plants. Two accessions were Afghani plants, and
they expressed three polymorphic sequences between them. One sequence was
shared by plants of Indian heritage, and the other two sequences were unique to
Afghani plants.
114 J.M. McPartland
Next-Gen sequencing (high-throughput sequencing) has generated a plethora of
genetic information. Van Bakel et al. (2011) used a whole genome shotgun
(WGS) method with Illumina technology to sequence Purple Kushand two hemp
cultivars, Finolaand USO-31.Van Bakel and colleagues also obtained tran-
scriptomes (cDNA libraries) from different tissues in these plants. Soon two other
Cannabis genomes were sequenced with WGS/Illumina machines, Chemdawg
and LA Condential(Medicinal Genomics Corporation 2011).
Tejkalová(2015) utilized Cannabis genomes (van Bakel et al. 2011) for
SNP-calling and genotyping with the KASP/SNPline platform. Haplotypes based
on 57 SNP positions for 44 samples of Sativaand 77 of Indicawere analyzed
with STRUCTURE. This probabilistic software identies the optimal number of
clusters (K) to divide a population, based on allele frequencies. Testing K values
from one to nine, the haplotype data best t K = 2 (two populations), but
STRUCTUREs assignment of individuals into Sativaand Indicamatched
poorly with their a priori identication.
Sawler et al. (2015) used genotyping-by-sequencing (GBS), which utilizes
restriction enzymes to break the genome into short reads (WGS uses random
ligation). They coupled ApeKI enzymes with Illumina machines for SNP discovery
and genotyping in ber-type and drug-type samples. GBS identied 14,031 SNPs
for analysis, after quality ltering. Drug-type strains were classied along a gradient
of ancestry proportions (percent Sativavs. percent Indica) reported in online
strain databases.
Their PCA analysis of genetic structure (SNP variations) using PLINK 1.9
clearly segregated 43 ber-type samples from 81 drug-type samples. The clusters of
Sativaand Indicapartially overlapped. Proportional ancestry in each sample
correlated moderately (r
0.36) with the principle component (PC axis 1) of
genetic structure. Similar results were obtained with fastSTRUCTURE, where data
from all 124 samples best t K = 2. The inability to separate Sativaand Indica
and the poor correlation of report ancestry was due, in part, to counterfeit strain
names: In a comparison of 17 paired samples with the same strain name, six pairs
(35%) were dissimilar, and shared more genetic similarity with other strain names.
Sawler calculated the xation index (F
) between subgroups based on
identity-by-state (IBS, implemented in PLINK). F
values range from 0 to 1; a
zero value indicates the subgroups interbreeding freely; a 1 value indicates the
subgroups are completely isolated from one another. The average F
ber- and drug-type plants was 0.156, which is similar to the degree of genetic
differentiation in humans between Europeans and East Asians. Average F
between ber-type plants and 100% Sativawas 0.161; F
between ber-type
plants and 100% Indicawas 0.136; no comparison was made between Sativa
and Indica.
Medicinal Genomics Corporation (2015) used Reduced Representation Shotgun
(RRS) sequencing to identify 100,000200,000 SNPs per strain. These data were
used to generate a nearest-neighbor tree with Purple Kush,”‘Finola,’‘USO-31,
and 50 ganjanym strains. Henry (2015) utilized open-access RRS data to evaluate
28 strains, using Adegenet 2.0. K-partition optimized at K = 1. PCA clustering with
4Cannabis sativa and Cannabis indica 115
a subset of 42 most-informative SNPs, however, clearly segregated three clusters:
Sativa(n = 17) Indica(n = 9), and two ber-type strains. These results were
conrmed with a neighbor-joining algorithm.
Lynch et al. (2015) sequenced 60 accessions using WGS, and added to this
dataset seven previous WGS reads (Van Bakel et al. 2011, Medicinal Genomics
Corporation 2011). For SNP-calling they aligned sequences with the draft genome
(Van Bakel et al. 2011). Then they sequenced 182 accessions using GBS, with
ECoRI and MseI restriction enzymes, for SNP-calling. A subset of 195 accessions
from WGS and GBS shared 2894 SNPs for analysis.
Two algorithms were used to K-partition the 195 accessions. FLOCK recognized
K = 3 groups, and fast STRUCTURE optimized the data at K = 2. The authors
went with FLOCK, because of perceived shortcomings in fast STRUCTURE,
although these perceived differences are contentious (Anderson and Barry 2015).
The K = 3 groups were recognized as WLD biotypes (e.g., Afghan Kush,
Chemdawg), NLD biotypes (e.g., Durban Poison,”“Easy Sativa), and a
polyphyletic hempgroup (e.g., Finola,’“AC/DC,Chinese hemp, Dagestan
Lynch and colleagues found no evidence for admixture (hybridization) in these
populations, based on results with the f
statistic and TreeMix. This seems unlikely,
given historical evidence of hybridization going back to the 1970s. TreeMix and the
statistic were developed with animal models; they may fall short with plants
having complicated histories of hybridization. TreeMix analyzes data with a
maximum of only 10 admixture (migration) events. The f
statistic must compare
three populations, so it was applied to the disputed FLOCK results.
They used vcftools to calculate F
between each FLOCK population.
= 0.099 between hempand combined NLD + WLD, and F
= 0.036
between WLDs and NLDs. More genetic heterozygosity existed within drug-type
plants (0.31%) than within ber-type plants (0.22%, signicant p< 0.001), which
they attributed to widespread hybridization of drug strainsan incongruous
hypothesis, given the previous paragraph.
Phylogenetic relationships between the 195 accessions were visualized in an
unrooted neighbor-joining networka phylogenetic tree with reticulation (diver-
gence and hybridization among ancestral lineages). The network revealed aspects of
ancestry not captured by a simple bifurcating tree, such as genetic admixtures
between Chinese hemp and feral hemp plants in the USA.
Next they pooled WGS data with GBS data from Sawler et al. (2015), with 4105
SNPs in common, and generated a neighbor-joining network with 210 accessions.
These data revealed a second NLD biotype clade, consisting of Indian, Southeast
Asian, and South African populations, along with various Hazehybrids. This
clade may represent accessions of Indian heritage with minimal admixture from
WLD biotypes. Lastly they pooled WGS data with both GBS datasets, a total of
289 accessions, ltered for overlapping SNPs (only 45 SNPs in commonthe two
GBS datasets were generated with different restriction enzymes), and used MEGA6
to generate a neighbor-joining tree.
116 J.M. McPartland
4.9 Conclusions
Research supports the classication of Sativaand Indica,but not their
nomenclature. Sativa(consistent with Lamarcksindica, the NLD biotype) differs
chemically and genetically from Indica(consistent with Vavilovsafghanica, the
WLD biotype). The systematics of these populations remains an open question.
Systematics adds the element of time to taxonomy. Sativaand Indicapre-
sumably diverged from a common ancestorbut when, and under what selection
pressures? One population evolved in low-warm-and-wet India, and the other in
high-cool-and-dry Afghanistan. Natural selection likely drove their initial
Good (1964) and Takhtajan (1986) divided the world into oristic regions
based on the distribution of distinctive (endemic) plant populations. The borders
between these oristic regions were delimited by natural barriers (geographic and
climatic) that prevented natural plant dispersal. Most of India lies in the Indian
Region. Afghanistan is part of the Irano-Turanian Region (Takhtajans term; Good
called it the Western and Central Asiatic Region). Another oristic region lies
between them, which includes most of Pakistanthe Sudano-Zambezian Region
(Takhtajans term), a.k.a. the North African-Indian Desert Region (Goods term).
Then humans took over with articial selection. In India, unpollinated females
were processed individually. Intentional selection of potent, high-THC individuals
was a straightforward process. In contrast, Afghani plants were processed in bulk,
with no selection of potent, high-THC individuals. Thus a millennium of selecting
different productsgañjāversus hashīshunintentionally drove divergence in
THC/CBD ratios. David Watson (pers. commun. 2012) stated that Afghani hashīsh
producers preferred certain terpenoids for aroma, and for physicochemical effects
on sifted hashīsh (e.g., the condensability of sesquiterpene alcohols). Consistent
with this, Hooper (1908) found the perceived quality and cost of three hashīsh
specimens from Kāšar correlated with their percentage of essential oil (i.e., ter-
penoids), and not with their percentage of resin (i.e., cannabinoids): Grade No. 1:
essential oil 12.7% and resin 40.2%; Grade No. 2: essential oil 12.4% and resin
40.9%; Grade No. 3: essential oil 12.0% and resin 48.1%.
Extensive cross-breeding between Sativaand Indicain the past 40 years has
rendered their distinctions almost meaningless in todays marketplace. Plants
should be identied by their chemical ngerprint, rather than characterizations such
as Sativa-dominant,”“Indica-dominant,or a whimsical strain name (Hazekamp
and Fischedick (2012); Hazekamp et al. 2016). Several analytical laboratories have
moved from cultivar to chemovar,and identify plants by their cannabinoid and
terpenoid content. These services include Strain Fingerprintby Steep Hill Labs,
PhytoFactsby Napro Research, Prole Testing by Werc Shop, and Know Your
Medicine by SC Labs.
However, as documented here, phytochemical and genetic research supports the
separation of NLD and WLD biotypes. Old landraces of Indian and Afghani
4Cannabis sativa and Cannabis indica 117
heritage face extinction through introgressive hybridization. We need to recognize
this biodiversity and conserve itfor future breeding efforts, at the very least.
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... They were based on illustrations by Anderson, which differed from the original botanical nomenclature. "Sativa" plants produce much more THC than CBD, while "Indica" plants produce almost equal amounts of THC and CBD, with a CBD/THC ratio of around 1 (McPartland, 2017). However, as McPartland also reported, these vernacular categories are unreliable for distinguishing between different chemotypes and/or cannabis end uses due to extensive cross-breeding and incomplete labelling during hybridisation (McPartland, 2017). ...
... "Sativa" plants produce much more THC than CBD, while "Indica" plants produce almost equal amounts of THC and CBD, with a CBD/THC ratio of around 1 (McPartland, 2017). However, as McPartland also reported, these vernacular categories are unreliable for distinguishing between different chemotypes and/or cannabis end uses due to extensive cross-breeding and incomplete labelling during hybridisation (McPartland, 2017). In addition, in most classification studies, samples had come from different sources and had been exposed to inconsistent environmental factors during growth phases, postharvest treatment, sample preparation, and extraction procedures during laboratory analysis (Jin et al., 2020(Jin et al., , 2021a. ...
... Only genetically female plants were cultivated, which were crossed and propagated with feminised seeds obtained by manipulating sex expression, as reported in our recent publication (Flajšman et al., 2021). This approach enabled us to develop two feminised high CBD breeding populations of medical cannabis, one corresponding to the "narrow leaflet drug type" (named MX-CBD-11) and the other one to the "wide leaflet drug type" (named MX-CBD-707) phenotype based on plant morphology according to McPartland (2017McPartland ( , 2018. ...
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Cannabis sativa L. is one of the oldest cultivated crops, used in medicine for millennia due to therapeutic characteristics of the phytocannabinoids it contains. Its medicinal properties are highly influenced by the chemotype, that is, the ratio of the two main cannabinoids cannabidiol (CBD) and Δ-9-tetrahydrocannabinol (THC). Based on published data, the chemotype should correlate with plant morphology, genetics, and photosynthetic properties. In this work, we investigated leaf morphology, plant growth characteristics, cannabinoid profiles, THCAS gene sequences, and plant photosynthetic traits in two breeding populations of medical cannabis (MX-CBD-11 and MX-CBD-707). The populations differed significantly in morphological traits. The MX-CBD-11 plants were taller, less branched, and their leaves had narrower leaflets than the bushier, wideleaved MX-CBD-707 plants, and there were significant differences between populations in the dry biomass of different plant parts. Based on these morphological differences, MX-CBD-11 was designated as a narrow leaflet drug type or vernacular “Sativa” type, while MX-CBD-707 was classified as wide leaflet drug type or “Indica” type. Chemical characterisation revealed a discrepancy between the expected chemotypes based on plant morphology; although both populations have high CBD, within each Type II (CBD/THC intermediate) and Type III (CBD dominant) plants were detected. The THCAS gene sequence analysis clustered the plants based on their chemotypes and showed high similarity to the THCAS sequences deposited in NCBI. In silico complementary analysis, using published molecular markers for chemotype determination, showed their low discrimination power in our two populations, demonstrating the genotype dependence of the molecular markers. Basic photosynthetic traits derived from light and CO 2 response curves were similar in the populations. However, measurements of gas exchange under chamber conditions revealed higher stomatal conductivity and photosynthesis in MX-CBD-707 plants, which were also characterised by higher day respiration. The results of this study showed that based on visual appearance and some morphological measurements, it is not possible to determine a plant’s chemotype. Visually homogenous plants had different cannabinoid profiles and, vice versa, morphologically distinct plants contained similar CBD and THC content. The two chemotypes identified in our experimental plants therefore did not correlate with plant visual appearance, leaf morphometry, and photosynthetic properties of the populations studied. Correlation was only demonstrated with the respect to THCAS sequences, which showed great discrimination power between the chemotypes.
... Cannabis is an annual dioecious plant belonging to the family Cannabaceae; dioecious means the male plants are distinct from the female plants. An early classification system distinguished C. sativa and Cannabis indica (McPartland, 2017). Their morphology (C. ...
... sativa originated from Europe and C. indica from Asia), and the composition of their secondary metabolites (secondary plant metabolites are large numbers of chemical compounds produced by plant cells, using biosynthetic enzymes and building blocks derived from primary plant metabolic pathways) all showed differences, but the dilemma about their taxonomical separation, whether they are separate species or subspecies, continues to the present day (Russo, 2007;Small, 2015;Pollio, 2016). Several review and research papers have provided detailed reports about the classification and nomenclature of cannabis (Fischedick et al., 2010;Pollio, 2016;McPartland, 2017McPartland, , 2018. The current consensus is that C. sativa and C. indica should not be considered different species. ...
... The high content of D 9 -THC classifies as drug-type cannabis (marijuana), whereas a low content is found in fiber-type cannabis (hemp) (Small and Beckstead, 1973). This fiber-type cannabis is characterized by a higher level of CBD, and the maximum D 9 -THC content of this type of cannabis was defined as 0.2%-0.3% of dry matter (McPartland, 2017). Chemotaxonomic evaluation has led to the recognition of an intermediate type with similar levels of D 9 -THC and CBD (Fischedick et al., 2010). ...
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Pain prevalence among adults in the United States has increased 25% over the past two decades, resulting in high health-care costs and impacts to patient quality of life. In the last 30 years, our understanding of pain circuits and (intra)cellular mechanisms has grown exponentially, but this understanding has not yet resulted in improved therapies. Options for pain management are limited. Many analgesics have poor efficacy and are accompanied by severe side effects such as addiction, resulting in a devastating opioid abuse and overdose epidemic. These problems have encouraged scientists to identify novel molecular targets and develop alternative pain therapeutics. Increasing preclinical and clinical evidence suggests that cannabis has several beneficial pharmacological activities, including pain relief. Cannabis sativa contains more than 500 chemical compounds, with two principle phytocannabinoids, Δ9-tetrahydrocannabinol (Δ9-THC) and cannabidiol (CBD). Beyond phytocannabinoids, more than 150 terpenes have been identified in different cannabis chemovars. Although the predominant cannabinoids, Δ9-THC and CBD, are thought to be the primary medicinal compounds, terpenes including the monoterpenes β-myrcene, α-pinene, limonene, and linalool, as well as the sesquiterpenes β-caryophyllene and α-humulene may contribute to many pharmacological properties of cannabis, including anti-inflammatory and antinociceptive effects. The aim of this review is to summarize our current knowledge about terpene compounds in cannabis and to analyze the available scientific evidence for a role of cannabis-derived terpenes in modern pain management. SIGNIFICANCE STATEMENT: Decades of research have improved our knowledge of cannabis polypharmacy and contributing phytochemicals, including terpenes. Reform of the legal status for cannabis possession and increased availability (medicinal and recreational) have resulted in cannabis use to combat the increasing prevalence of pain and may help to address the opioid crisis. Better understanding of the pharmacological effects of cannabis and its active components, including terpenes, may assist in identifying new therapeutic approaches and optimizing the use of cannabis and/or terpenes as analgesic agents.
... There are a lot of debates on the taxonomic interpretations in which some researchers used the term 'hemp' referring to Cannabis sativa and 'marijuana' referring to Cannabis indica although it has also been used to refer to other fibre crops (Clarke & Watson, 2007;McPartland, 2017). Cannabis sativa may primarily be grown for three purposes namely industrial, recreational, and medicinal (Farinon et al., 2020;Rupasinghe et al., 2020). ...
There is an abundance of international studies that have lauded the benefits of medicinal cannabis for various illnesses. Likewise, local studies on cannabis in Malaysia that initially revolve around the harmful effects of cannabis misuse are increasingly shifting towards the benefits of medicinal use. Pursuant to scientific discovery of the medicinal benefits of cannabis, a myriad of literature begins to focus on the legality of utilising cannabis for medicinal purposes. Whilst many countries around the world, some of those including South East Asia countries have legalised medicinal cannabis, Malaysia is still lagging behind. As of now, there is no local study focusing on the legalisation of medicinal cannabis, leaving a gap in providing a reference to Malaysia's way forward on this issue. Under the current legislative regime, the use of cannabis is prohibited under the Dangerous Drugs Act (DDA) 1952 [Act 234]. However, the recent case of Muhammad Lukman Mohamad has sparked national interest that triggered calls for decriminalisation and legalisation of cannabis for medical reasons. Following these calls, it is pertinent to examine and critically analyse the medical, ethical, and legal considerations on the legalisation of medicinal cannabis in Malaysia. Such consideration is deemed necessary to protect the health and safety of patients. This research adopts an exploratory research design and employs a qualitative research approach in examining medical, ethical, legal, and theological considerations towards the legalisation of medicinal cannabis. Comparative legal analysis covers substantive and procedural components of jurisdictions in selected countries that have legalised medicinal cannabis such as the United Kingdom (UK), Canada, and Thailand. Examination of medico-ethical considerations is done using Biomedical Ethics Theory and Philosophical Theory. These considerations provide a comprehensive analysis of the legalisation of medicinal cannabis in Malaysia that will contribute to the current body of knowledge. This research proceeds to make several recommendations deemed relevant towards the legalisation of medicinal cannabis in Malaysia.
... The WLD biotype generally produces much greater amounts (relative to the weight of the plant) of the major cannabinoids, especially THC and CBD, mainly because of its denser flowers and greater expression of glandular trichomes. On the other hand, the NLD biotype produces larger amounts of the minor cannabinoids, such as THCV (McPartland 2017). The THC to CBD ratio often favours CBD in WLD type (especially native Afghan) plants, but this is very unusual in the NLD type which often produces only trace amounts of CBD . ...
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Cannabis spp. are some of the most controversial medicinal plants in the world. They contain great amounts of biologically active secondary metabolites, including the typical phenolic compounds called cannabinoids. Because of their low toxicity and complex biological activities, cannabinoids can be useful in the therapy of various diseases, but adverse psychological effects (of Δ9-THC in particular) raise concerns. This review summarizes the current knowledge of selected active C. indica compounds and their therapeutic potential. We summarize the main compounds contained in cannabis, the mechanisms of their effects, and their potential therapeutic applications. Further, we mention some of the clinical tests used to evaluate the efficacy of cannabinoids in therapy.
In the space of two years, selling cannabis in Canada went from being a criminalized activity to an essential service. However, the broad recognition of its relative safety compared with other intoxicants and potential medical benefits are outpacing regulatory strategies to manage its responsible use. The paradoxical normalization of cannabis is based on its contested legal, moral, and cultural status. This paper makes three contributions. First, based on secondary data analysis, we present qualitative data from cannabis insiders in the province of British Columbia (B.C.), focused on their perceptions of cannabis as medicine, consumer good, and nuisance crime. Second, we consider tolerance and aversion related to cannabis normalization by linking the perceptions of these insiders with past and recent research. Finally, we argue that the lack of legal spaces to consume cannabis exacerbates concerns about impaired driving, nuisance crime, and stigmatization. Applying levels of stigma to areas of cannabis liberalization can assist efforts to describe the paradoxes of normalization. These appear to result from older protection-based policies, which may have the effect of prolonging what they seek to disrupt.
Marijuana (Cannabis) belongs to family Cannabaceae. This plant is a great natural source of cannabinoids, which have several medical applications such as it is used for the treatment of anxiety and psychosis, disorders of motility and epilepsy. However, this plant is also notorious for its pollen allergic reactions. Mainly two species, that is, C. sativa L. and C.indica Lam. are involved in allergic reactions. In this study, morpho‐palynological studies of both the species were conducted. A number of morphological differences were observed in the both species. C. sativa is tall with narrowed leaves and fibrous stalk plant while C. indica is short, bushier, broad leaves, and having woody stalk plant. Both the species are varied in their pollen characteristics. The polar diameter of C. sativa observed was 7.5 μm and equatorial diameter was 8 μm, and its shape was oblate‐spheroidal. C. indica had 7 μm polar diameter, 7.5 μm equatorial diameter but its shape was similar to C. sativa, that is, oblate‐spheroidal. About 40%‑50% people were affected with both these marijuana species and had symptoms of allergies like dry cough, congestion, itchy eyes, sore throat, nausea, a runny nose, watery eyes, and sneezing. It was concluded that both species' pollen are allergy causing and pollen of both species are different with variations in morphological and allergic symptoms. Marijuana has great therapeutic importance but it is also notorious for its pollen allergic reactions. Two species of Marijuana, that is, Cannabis sativa L. and Cannabis indica Lam mainly responsible for these pollen allergies. The present research work focused the morpho palynological analysis of these two Marijuana species through light and scanning electron microscopy. Palynological analysis of Marijuana species causing allergies.
Cannabis is the most widely used recreational drug in the world. Cannabis sativa and Cannabis indica have been selectively bred to develop their psycho‐active properties. The increasing use in many countries has been accelerated by the COVID‐19 pandemic. Cannabis can provoke both type 1 and type 4 allergic reactions. Officially recognised allergens include a pathogenesis‐related class 10 allergen, profilin and a non‐specific Lipid Transfer Protein. Other allergens may also be relevant, and recognition of allergens may vary between countries and continents. Cannabis also has the potential to provoke allergic cross‐reactions to plant foods. Since cannabis is an illegal substance in many countries, research has been hampered, leading to challenges in diagnosis since no commercial extracts are available for testing. Even in countries such as Canada, where cannabis is legalized, diagnosis may rely solely on the purchase of cannabis for prick‐to‐prick skin tests. Management consists of avoidance, with legal issues hindering the development of other treatments such as immunotherapy. Education of healthcare professionals is similarly lacking. This review aims to summarise the current status of cannabis allergy and proposes recommendations for the future management of this global issue.
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Marijuana and hemp (Cannabis) and the closely related hop genus (Humulus) are the only widely known genera included in the small, but economically valuable, Cannabaceae family. Swedish botanist Carl Linnaeus, the “father of modern taxonomy,” first published the scientific name Cannabis sativa in his seminal Species Plantarum of 1753. The Latin name Cannabis derives from Greek (kannabis) and may have been originally derived from Scythian. The term sativa simply means “cultivated” and describes the common hemp plant that was widely grown across Europe in Linnaeus’ time. We, the authors, consider C. sativa to be native to western Eurasia and especially Europe, where, for millennia, the plant has been grown for its strong fibers and nutritious seeds, and from where it was introduced to the New World multiple times during early European colonization. Cannabis sativa plants also produce very small amounts of the compound delta-9-tetrahydrocannabinol (THC), the medically valuable and primary psychoactive cannabinoid found only in Cannabis. Since C. sativa evolved within the geographical limits of western Eurasia, it represents only a small portion of the genetic diversity seen in the genus Cannabis worldwide.
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Cannabis: Evolution and Ethnobotany is a comprehensive, interdisciplinary exploration of the natural origins and early evolution of this famous plant, highlighting its historic role in the development of human societies. Cannabis has long been prized for the strong and durable fiber in its stalks, its edible and oil-rich seeds, and the psychoactive and medicinal compounds produced by its female flowers. The culturally valuable and often irreplaceable goods derived from cannabis deeply influenced the commercial, medical, ritual, and religious practices of cultures throughout the ages, and human desire for these commodities directed the evolution of the plant toward its contemporary varieties. As interest in cannabis grows and public debate over its many uses rises, this book will help us understand why humanity continues to rely on this plant and adapts it to suit our needs.
The relationship between glandular trichomes and cannabinoid content in Cannabis sativa L. was investigated. Three strains of Cannabis, which are annuals, were selected for either a drug, a non-drug, or a fiber trait and then cloned to provide genetically uniform material for analyses over several years. The distribution of the number and type of glands was determined for several organs of different ages including the bract and its subtending monoleaflet leaf and the compound leaf on pistillate plants. Quantitation of glands on these structures was integrated with gas chromatographic analyses of organ cannabinoid profiles. A negative correlation was found between cannabinoid content and gland number for each of the three clones. Isolated heads of the capitate-stalked glands also were analyzed for cannabinoid content and found to vary in relation to clone and gland age. These studies indicate that cannabinoids may occur in plant cells other than glandular trichomes. The results of these studies emphasize the need for stringent sampling procedures in micromorphological studies on trichome distribution and analytical determinations of cannabinoid content in Cannabis.
Trichome density and type and cannabinoid content of leaves and bracts were quantitated during organ ontogeny for three clones of Cannabis sativa L. Trichome initiation and development were found to occur throughout leaf and bract ontogeny. On leaves, bulbous glands were more abundant than capitate-sessile glands for all clones, although differences in density for each gland type were evident between clones. On pistillate bracts, capitate-sessile glands were more abundant than the bulbous form on all clones, and both types decreased in relative density during bract ontogeny for each clone. The capitate-stalked gland, present on bracts but absent from vegetative leaves, increased in density during bract ontogeny. The capitate-stalked gland appeared to be initiated later than bulbous or capitate-sessile glands during bract development and on one clone it was first found midway in bract ontogeny. Nonglandular trichomes decreased in density during organ ontogeny, but the densities differed between leaves and bracts and also between clones. Specific regulatory mechanisms appear to exist to control the development of each trichome type independently. In addition, control of trichome density seems to be related to the plant organ and clone on which the gland type is located. Cannabinoid synthesis occurs throughout organ development and is selectively regulated in each organ. Typically, cannabinoid synthesis occurred at an increasing rate during bract development, whereas in developing leaves synthesis occurred at a decreasing rate. Cannabinoid content on a dry weight basis was generally greater for bracts than leaves. Analyses of leaves indicate that other tissues in addition to glands may contain cannabinoids, while for bracts the gland population can accommodate the cannabinoid content for this organ. The functional significance of trichomes and cannabinoids in relation to evolution is discussed.
Wild growing Cannabis from different altitudes and locations was collected in northern India. Plant material from each collection was analyzed quantitatively for 10 cannabinoids by GC. The average total cannabinoid content of plants growing above 2,000 m was 1.33%, whereas below 2,000 m the average total cannabinoid content was 2.74%. Morphological characteristics were recorded. Data obtained from wild growing Indian Cannabis (20 variants) are compared with data obtained from the same variants grown in Mississippi, USA.