overlap and outliers.“Indica”strains produced more guaiol,c-eudesmol, andb-
eudesmol, as well as another sesquiterpene alcohol—a-bisabolol, plus three mon-
terpene alcohols: a-terpineol,b-fenchol, and linalool. “Sativa” strains 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, “Indica” samples 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“Indica”strains. They also report significantly more limo-
nene and myrcene in“Indica,”consistent with Hillig’s 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
significant amounts ofb-eudesmol andc-eudesmol, like plants of Afghani heritage.
Casano et al. ( 2011 ) compared 16 unnamed hybrid accessions, characterized as
“mostly Indica”or“mostly Sativa.”“Mostly Indica”plants produced significantly
higher levels of limonene,b-myrcene, and camphene.“Mostly Sativa”produced
significantly higher levels of sabinene,Δ-3-carene,a-phellandrene, 1,8-cineole, cis-
b-ocimene, trans-b-ocimene, anda-terpinolene.
Elzinga et al. ( 2015 ) assigned strains to“Sativa”or“Indica”according to the
Leafly database, as described earlier. They noted that strains namedKush,“char-
acteristic of the wide leaflet drug type strains originating from Hindus Kush region
of Afghanistan and Pakistan,”contained higher levels of guaiol,b-eudesmol,b-
myrcene, trans-ocimene, andb-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 ofb-myrcene
anda-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 significant 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 thatb-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-lock”of“Indica”tob-myrcene, and Russo ( 2016 ) attributed the uplifting
effects of“Sativa”to 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”
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