(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 “Indica” and “Sativa”
(Elzinga et al. 2015 ; Lynch et al. 2015 ).
Regardingb-myrcene, earlier studies showed greater amounts in Afghani lan-
draces than in plants of Indian heritage, albeit short of statistical significance (Hood
and Barry 1978 ; Hillig 2004b). This trend continued in four recent studies of
“Indica”versus“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.
2015 ).
4.8 Genetic Studies
Hillig (2005a) analyzed allozyme variation in the sameCannabiscollection 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 andfiber-type plants into distinct clusters, but the WLD ellipse and
NLD ellipse substantially overlapped.
Gilmore et al. ( 2007 ) examined 76Cannabisaccessions forfive polymorphic loci
sequenced from chloroplast and mitochondrial DNA. The study’sflaws are manifold,
but parsimony analysis recovered three clades. Clade A comprised a majority of
fiber-type plants. Clade B included Afghani landraces along with most drug strains—
hybrids and police seizures. Clade C was the most interesting—nothing but classic
“Sativas”: 12 landraces from India, Nepal, Thailand, Jamaica, Mexico, and Africa.
Gilmore ( 2005 ) gave the nameC. sativa rastato plants in Clade C.
Knight et al. ( 2010 ) tested six seized plants, identified by their morphology as
“Sativa”(n = 2) or“Indica”(n = 4). Five short tandem repeat (STR) loci, analyzed
with PCA, clearly segregated“Sativa”plants from three of the“Indica”plants. The
fourth“Indica”exhibited a unique genotype suggestive of a polyploid condition.
Piluzza et al. ( 2013 ) compared 19 accessions: one Afghani,five of Indian her-
itage, three“Skunk”hybrids, and an assortment offiber-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 offiber-type plants. The cluster of Indian heritage plants
was sister to the“Skunk”cluster.
Onofri et al. ( 2015 ) searched for single nucleotide polymorphisms (SNPs) in
THCA-S sequences. They found nine unique THCA-S sequences amongst 18
accessions offiber- 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