Gland Distribution and Cannabinoid Content in Clones of Cannabis sativa L.

Author(s): Jocelyn C. Turner, John K. Hemphill and Paul G. Mahlberg

Source: American Journal of Botany, Vol. 64, No. 6 (Jul., 1977), pp. 687-693
Published by: Botanical Society of America
Stable URL: http://www.jstor.org/stable/2441721 .
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Amer. J. Bot. 64(6): 687-693. 1977.

GLAND DISTRIBUTION AND CANNABINOID CONTENT

IN CLONES OF CANNABIS SATIVA L.'
JOCELYN TURNER, JOHN K. HEMPHILL,
C. AND
PAUL G. MAHLBERG
Departmentof Plant Sciences,Indiana University,
Bloomington47401

A B S T R A C T
The relationship betweenglandulartrichomes and cannabinoidcontentin CannabissativaL.
was investigated. Threestrainsof Cannabis,whichare annuals,wereselectedforeithera drug,
a non-drug, or a fibertraitand thenclonedto providegenetically uniformmaterialforanalyses
over severalyears. The distribution of thenumberand typeof glandswas determined forsev-
eral organsof different ages includingthe bract and its subtendingmonoleafletleaf and the
compoundleaf on pistillateplants. Quantitationof glandson thesestructures was integrated
withgas chromatographic analysesof organcannabinoidprofiles.A negativecorrelationwas
foundbetweencannabinoidcontentand gland numberforeach of the threeclones. Isolated
heads of the capitate-stalked glandsalso were analyzedforcannabinoidcontentand foundto
varyin relationto clone and glandage. These studiesindicatethatcannabinoidsmay occur in
plantcells otherthanglandulartrichomes.The resultsof thesestudiesemphasizethe need for
stringentsamplingproceduresin micromorphological studies on trichomedistributionand
analyticaldeterminations of cannabinoidcontentin Cannabis.

GLANDULAR trichomescoveringthe plant surface The purposeof thisinvestigation is to determine

have been implicatedas the source of cannabi-
noids in Cannabis sativa L. (DePasquale, 1974;
Malingre et al., 1975). Hammond and Mahl-
berg (1973), in their scanning electron micro-
scope study of these trichomes,described three
gland types: the bulbous gland which possesses
a veryshortstalk and a small head, the capitate-
sessileglandwhichhas a largeglobularand multi-
cellular head, and the capitate-stalkedgland
which consistsof a large multicellularhead that
terminatesa stalk of variable height. Nongland-
ular trichomesare also presentin abundance on
the plant epidermis (Ledbetter and Krikorian,
1975). Fairbairn (1972) reportedthe presence
of cannabinoids in both capitate-sessile and
capitate-stalkedglandsand indicatedthatcapitate-
stalked glands were the major cannabinoidcon-
tainingglands. DePasquale (1974), in an ultra-
structuralstudy of the capitate-stalkedglands,
interpretedthe secretorypart of the gland to be
a combinationof the gland head and perhaps
apical stalk cells. Malingre et al. (1975) con-
cluded that cannabinoidswere presentmainlyin
the epidermal glands and not in the mesophyll
cells or nonglandulartrichomes,although there
was some indicationof cannabinoid content in
theleafmidrib.
whethera correlationexistsbetweenglandulartri-
chomes and cannabinoidcontentin Cannabis. If
specific glands are associated with cannabinoid
content,a correlationshould exist between the
gland numberpresenton a particularplant part
and the cannabinoidcontentof that part. Fur-
thermore,it should also be possible to establisha
correlationbetween the numbersof each gland
typeand cannabinoidcontent.
MATERIALS AND METHODS-Clones-Plants of
three Cannabis strains,which are typicallyan-
nuals, were cloned to providematerialof genetic
homogeneity.These strainsincluded a high A9-
tetrahydrocannabinol (A9-THC) strain (152), a
low A9 THC strain(79), and a fiberstrain(87).
Plants were grown in a loam-vermiculite-sand
mixture(6:2:1) under ambientgreenhousecon-
ditions. Cuttingswere taken fromthe parentpis-
tillateplant of each strain,treatedwithRootone,
and rootedin perlite. Three-week-oldrootedcut-
tingswere transplantedinto the above soil mix-
ture. Cuttingsderivedfromvegetativeor flower-
ing plants were made at regularintervalsfrom
the parentor previouscuttingsto providea con-
tinuouspopulationof each clone throughoutthe
year.

Plant partssampled-Selected plantpartsfrom

1 Received for publication 22 September 1976; re-
visionaccepted26 February1977. each strainwere analyzed for theircannabinoid
This researchwas supportedby a grantfromthe Na- contentand glandnumber.Young pistillatebracts
tional Instituteon Drug Abuse (DA 00981) to P. G. were approximately 3 mm in lengthand enclosed
Mahlberg. The authorsthankDr. ErnestSmall for the an unpollinatedflower. Mature bracts were ap-
seed used to establishthe clones employedin this in-
No. P10043113(PGM). proximately
vestigation.D. E. A. Registration 7 mm in lengthand enclosed a seed.
687

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688 AMERICAN JOURNAL OF BOTANY [Vol. 64

TABLE 1. Quantitationof glands on bracts acetate. Samples were driedby the criticalpoint
techniqueusing CO2 (Anderson, 1951; Horridge
Glands/mm2 and Tamm, 1969), mountedon stubswithepoxy
Bulbous and glue, coated with a layer of carbon followedby
immature
Bract Capitate- Capitate- capitate- gold-palladiumby utilizing a Rotilt shadower
Clone age stalked sessile stalked* (EBTEC Corp), and subsequentlyexaminedwith
152 Young 0.6 72.0 48.6 an ETEC Autoscan SEM operatedat 20 kV.
(drug) Mature 34.2 41.1 8.2
Gland quantitation-Gland number per unit
79 Young 10.4 66.1 83.5 area was determinedby countingglands directly
(non-drug) Mature 17.1 32.8 10.4
on the SEM screen. At a magnification of 360,X
87 Young 0 149.3 157.3 witha tiltof 250, the observedfield was '/ mm
(fiber) Mature 37.3 10.0 0.7 on a side (1/16 mm2). All glandular and non-
*
Bulbous and immaturecapitate-stalked glands were glandulartrichomeswithinthe fieldwere counted.
in dis-
scored as one categorybecause of the difficulty For both the young and maturebracts, random
tinguishingbetweenthe two typesat a young stage in fieldstotaling2 sq mm were counted. On floral
development. leaves, countswere made at the midpointof the
leaf blade fromthe midribto the margin. Multi-
ple counts (16 fields,totaling1 sq mm) of both
Floral leaves, generallymonoleafletleaves, sub- adaxial and abaxial vein and nonveinareas were
tendingyoung and maturepistillatebracts were made, and the resultswere averagedto provide a
analyzed as were maturecompound leaves from mean forthe sample. On compoundleaves, gland
vegetativeregionsoftheaxis. countswere made at the tip, midpoint,and base
of the leaf along the midriband midwaybetween
Gas chromatography-Plantparts to be ana- the midriband the leaf margin. The numberand
lyzed were collectedat a giventime (3 P.M.) and type of fields counted were the same as for the
immediatelyoven dried at 60 C for 12 h. floralleaves. For all samples,the standarddevia-
Weighed samples were extractedin covered test tion was determinedand criticalvalues of t were
tubes with spectrogradechloroform(3X) for at calculated to test for significancewhen means
least 1 h/extractionat 4 C. Plant materialwas werecompared.
removedby filtrationand the combined aliquots
were concentratedto drynessby gentle nitrogen Single gland sample-Cannabinoid contentof
evaporation. Cannabinoids were dissolved in 1 single capitate-stalkedglands was determinedby
ml ethanol containingthe internalstandard Ai4- removingglands witha tungstenneedle and ana-
androstene-3,17-dione(1 mg/ml;w/v) (Dooren- lyzingthemby gas chromatography as previously
bos et al., 1971). A 0.5 ,u sample of the testsolu- described. Only intact gland heads were taken
tion was injectedinto the chromatograph. for the sample. Samples consistedof 20 gland
Analyseswereperformed on a Hewlett-Packard heads, althoughfor two samples an additional
5710A chromatographequipped with hydrogen 100 gland heads were taken to provide a greater
flame-ionization detectorsand programmedfrom concentrationof extractablematerial for ascer-
180 to 240 C at 4 C/min with an additional 8 tainingthe cannabinoidspectrumof glands.
min isothermalperiod (240 C). The inlet tem-
peraturewas 250 C and the detectortemperature RESULTS-Gland qulantitation of bracts-The
was 300 C. Glass columns (2 mm,id x 2.43 m) numberof capitate-stalkedglands per unit area
were treatedwith 5% dimethyldichlorosilane in of the abaxial surfaceof the bract increasedwith
toluene and packed with 3% OV-1 on 80/100 maturity of the bractin all threestrains(Table 1;
mesh Supelcoport. The head pressuresfornitro- Fig. 1, 2). Mature bracts of clones 152 and 87
gen, hydrogen,and compressedair were 50, 15, had a similarnumber of capitate-stalkedglands
and 40 lbs/sq in, respectively.Peak area of each (34 and 37 glands/mm2,respectively) while
cannabinoidwas comparedwiththe peak area of clone 79 had only half that number(17 glands/
theinternalstandardforcannabinoidquantitation. mm2). Capitate-sessileglands were abundanton
Data in tables are mean values of threereplicates. theyoungbractsof all strainswithtwicethenum-
ber (149 glands/mm2)presenton clone 87 than
Scanningelectronmicroscopy-Collections of on clones 152 and 79. The numberof capitate-
plant organswere made at a giventime (3 pm), sessileglandsper unitarea decreasedduringbract
fixed 18-20 h in 4% glutaraldehydeand 0.7% enlargementand fewer capitate-sessile glands
picric acid in 0.1 M sodium cacodylate (Turner, (10/mM2) were presenton the maturebracts of
1970), washed in 0.1 M sodiumcacodylateat pH clone 87 than on clones 152 (41 glands/mm2)
7.2, post-fixedin 1% OS04 in sodiumcacodylate and 79 (32 glands/mm2). Clones also varied in
for 1 h, rinsed with water, and dehydrated the numberof bulbous glands per unit area on
througha graded series of ethanols into amyl maturebracts: eightto ten glands were present

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July, 1977] TURNER ET AL.-CANNABINOID CONTENT OF CANNABIS 689

11

gland; T, nonglandulartri-
gland; St, capitate-stalked
KEY TO LABELING: B, bulbous gland; Se, capitate-sessile
chome; SO, stomate,V, vein.
Fig. 1, 2. Young and maturebracts. 1. Young bractsshowingabundantbulbous and capitate-sessile glands
trichomes.X 70. 2. Maturebractshowingpresenceof capitate-stalked
as well as non-glandular glandsin addition
to nonglandulartrichomes.x 70.

on clones 152 and 79 whileless thanone per unit
area occurredon clone 87. The ratioof the three
gland typeson maturebractsalso variedbetween
clones. In bothyoungand maturebracts,capitate-
sessile glands occurred in greaternumbersthan
capitate-stalkedand bulbous glandson clones 152
and 79, but on maturebractsof clone 87 capitate-
stalkedglands predominatedover sessile glands,
whilebulbous glands were fewin number.
Cannabinoidcontentof bracts-Determination
of cannabinoid content in young and mature
bracts,which correspondedto samples analyzed
forglanddistribution,revealedthatonlyclone 87
had an increase in all cannabinoidsinvestigated
as thebractmatured(Table 2). Clones 152 and
79 both showed a decrease in cannabichromene
(CBC) and A9-THC contentwith maturationof
the bract. However, 152 showed an increase in
cannabidiol(CBD) and cannabinol (CBN) con-
tentwithmaturationwhileclone 79 showed a de-
crease. Relatively low concentrationsof CBC
and CBN were foundin bracts of all strains. A
positive correlationbetween the concentrations
of CBC and CBN withthe characteristiccanna-
binoid of each strainwas observedwith the ex-
ceptionofCBN in clone 152.
Gland quantitationof floral leaves-Capitate-
stalked glands were present on mature floral
leaves of all threeclones with 87 having a con-
siderablyhighernumberper unitarea (11 glands/
mm2) than eitherclone 152 or 79 (Table 3).
Capitate-stalkedglands increased with leaf ma-
turityin clone 79, but decreased in clone 152.
Capitate-sessileglands increasedwith leaf matu-
rityin clone 152 fromnine to 11 glands/mm2,
TABLE 3. Quantitation
of glands on floralleaves

Glands/mm2
TABLE 2. Cannabinoidcontentof bracts
Leaf Capitate- Capitate-
Clone age stalked sessile Bulbous
mg Cannabinoids/100mg drywgt
Bract 152 Young 0.6 9.5 18.8
Clone age CBD CBC A9-THC CBN
(drug) Mature 0.2 11.7 6.2
152 Young 0.02 0.45 6.64 0.17 79 Young 0 6.5 23.7
(drug) Mature 0.10 0.10 5.41 0.39 (non-drug) Mature 2.3 2.7 13.0
79 Young 5.38 0.21 0.22 0.11 87 Young* - -
(non-drug)Mature 3.02 0.12 0.18 Trace (fiber) Mature 11.0 8.3 16.8
87 Young 3.66 0.16 0.20 Trace * Nonglandular trichome density obstructed view of
(fiber) Mature 5.33 0.33 0.28 0.28 the leaf surface and prevented gland quantitation.

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690 AMERICAN JOURNAL OF BOTANY [Vol. 64

TABLE 4. Cannabinoid content of floral leaves Observationsof the midpointof leaves fromclone
152 (Table 5) showed that adaxial and abaxial
Leaf
mg Cannabinoids/100mg dry wgt surfacesof the leaves as well as vein and non-
Clone age CBD CBC A9-THC CBN vein regionswere significantly in gland
different
152
distribution(Fig. 3-6). This patternwas found
Young 0.07 0.68 4.03 0.19
(drug) Mature Trace Trace 1.92 0.12
in all six sample areas. No capitate-stalkedglands
were present. For quantitativecomparison of
79 Young 5.00 0.20 0.22 0.32 capitate-sessileand bulbous glands, a mean was
(non-drug) Mature 1.79 0.11 0.11 Trace calculated for each sample area (Tables 5, 6).
87 Young 3.24 0.16 0.16 0.10 Data evalulatedstatistically by means of the t test
(fiber) Mature 4.30 0.24 0.94 0.16 showed no significantdifferenceamong the six
areas. Clones 87 and 79 werefoundto followthis
sampling pattern as described for clone 152.
Therefore,data from all six areas on the leaf
but decreasedin clone 79 fromsix to two glands/ were combined to give a mean for each gland
mm2. Bulbous glands decreased with leaf ma- typeobservedon compoundleaves in each clone
turityin both 152 and 79. High densitiesof tri- (Table 7).
chomes of young floral leaves of clone 87 ob- Gland typeand numbervariedwiththe respec-
structedtheviewof theleaf surfaceand prevented tive clones. Clone 79 had significantly more cap-
gland quantitation. As in the bracts, the rela- itate-sessileand bulbous glands/mm2than either
tive numbersof glands varied among the clones. 152 or 87. Clone 152 had the lowestnumberof
On the mature floral leaves, clone 87 had the bulbous glands (4.5/mm2), but clone 87 had the
highestnumber of capitate-stalkedand bulbous lowest number of capitate-sessileglands (2.6/
glands, while 152 had the highest number of mm2). Gland number and type on compound
capitate-sessile
glands. leaves varied considerablyas comparedto bracts
and floralleaves (Tables 1, 3, 7).
Cannabinoid content of floral leaves-Com-
parisonsof cannabinoidlevels in leaves subtend- Cannabinoid contentof compound leaves-In
ing young and maturebracts (Table 4) showed general,cannabinoidcontentof compoundleaves
thatclones 79 and' 87 followedthe same patterns (Table 8) was lower thanthatforbracts (Table
as foundin the bracts (Table 2). Cannabinoids 2) and floralleaves (Table 4). However,as ex-
in clone 79 decreased as the leaf matured,while pected,compoundleaves of clone 152 contained
theyincreasedin clone 87. In contrast,the can- a higherlevel of A9-THC than eitherof the other
nabinoid content in clone 152 decreased with two clones. Clone 87 containedthe highestlevel
leaf maturation(Table 4), whereas in the bract of CBD as well as the highesttotal cannabinoids.
some cannabinoidsincreased in quantityas the Clone 79 had approximately
bracts matured (Table 2). All three clones the same concentra-
tion of A9-THC as clone 87. However,the CBD
showedthe same quantitativetrendof the charac-
teristiccannabinoidas well as the principalcan- contentwas lower in clone 79 than clone 87.
nabinoids with maturationof both bracts and Only trace amountsof CBC and CBN were de-
floralleaves (Table 2, 4). tectedin all strains.

Gland quantitationof compound leaves-The Cannabinoid contentof single capitate-stalked

large size of compoundleaves made it necessary glands-Three different stages of maturationof
to establisha samplingprocedurefor quantitat- capitate-stalkedglands could readily be recog-
ing the distributionof glands on the leaf surface. nized withina populationof these glands on ma-
Six areas of the leaf were chosen for sampling. ture bracts. Initially,the maturegland possessed

TABLE 5. Quantitation of glands at midpoint of compound leaves of clone 152

Glands/mm2

Midrib Betweenmidriband leaf margin

Leaf surface Region Capitate-sessile Bulbous Capitate-sessile Bulbous
Adaxial Vein 0 0 0 0
Nonvein 1.6 0 2.2 0
Abaxial Vein 1.6 3.9 0.7 0
Nonvein 14.9 11.4 12.8 13.4
Samplemean 5.5 4.5 4.3 3.6

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July, 1977] TURNER ET AL.-CANNABINOID CONTENT OF CANNABIS 691

Fig. 3-6. Regionsof leaf surface. 3. Adaxial vein region showingabundantnonglandulartrichomes,a few
glands. X 430. 4. Adaxial nonveinregionwithfewnonglandular
bulbousglands,and no capitate-sessile trichomes.
glands are sometimespresentas well. X 430. 5. Abaxial vein regionshowingabun-
Bulbous and capitate-sessile
dantnonglandular trichomesand a few bulbousand capitate-sessileglands. x 430. 6. Abaxial non-veinregion
withnumerousnonglandular trichomes, glands, and abundantstomata. x 430.
a fewbulbousand capitate-sessile
See page 691 for KEY TO LABELING.

a head witha clearliquidcontent.The second istic of the strain (CBD, 229 ng/gland) than
stagewas representedby the aged glandwhich did the glands fromclone 152 (A9-THC, 57 ng/
appearedyellowand containeda sticky,more gland). Withaging,clone 87 regularlycontained
solidcontentin the head, whilethe thirdstage higherlevels of its characteristic
cannabinoidthan
by a senescent
was represented glandwitha red, clone 152. During the progressivematurationof
driedhead. Cannabinoidanalysesof thesethree glands in clone 152, the CBN contentaccumu-
glandpopulationson clones 152 and 87 (Table lated in the glands and then decreased sharply
thatthecannabinoid
9) indicated contentof the (Table 9). CBC was not detectedin the glands
glandsdecreasedsignificantly
capitate-stalked as of eitherstrain.
mature
theglandheadaged. In addition, capitate-
stalkedglandsfromclone 87 containedhigher DISCUSSION-Floral and vegetativeorgans of
concentrationsof the cannabinoidcharacter-Cannabis have been previouslyanalyzedfor pos-

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692 AMERICAN JOURNAL OF BOTANY [Vol. 64

TABLE 6. Quantitationof glands on compoundleaves TABLE 8. Cannabinoidcontentof compoundleaves

of clone152
mg Cannabinoids/100mg drywgt
Glands/mm2 Clone CBD CBC A9-THC CBN
Betweenmidrib
Midrib and leaf margin 152 (drug) Trace Trace 0.37 0.04
Leaf Capitate- Capitate-
regions sessile Bulbous sessile Bulbous 79 (non-drug) 1.49 0.06 0.06

Tip 5.3 4.3 5.2 6.3 87 (fiber) 2.38 0.07 0.08 Trace

Midpoint 5.5 4.5 4.3 3.6

Base 2.9 4.6 5.1
sible sites of cannabinoidsynthesis(DePasquale,
1974; Crombie and Crombie, 1975; Malingre et
al., 1975). If as postulated,cannabinoids are
synthesizedin the glandular trichomeson the
plantsurface,thengland numbershould correlate
withcannabinoidcontent.In our attemptto dem-
onstratethispossiblecorrelation,certainsampling
difficultieswere encountered.Differentplant or-
gans vary in theircannabinoidcontent(Dooren-
bos et al., 1971; Fettermanet al., 1971) and
gland type (Fairbairn, 1972; Hammond and
Mahlberg,1973; Ledbetterand Krikorian,1975;
Dayanandan and Kaufman,1976). Therefore,an
attemptwas made to standardizeboth quantita-
tive and qualitativeanalysesfortheseparameters.
Since cannabinoid profileshave been shown to
differamong Cannabis variants(Small, 1973) as
well as withina singlestrain(Small, 1973; Coff-
man and Gentner,1975; Latta and Eaton, 1975),
threestrainswere selectedthatpossessed eithera
drug,non-drug,or fibertraitand then cloned to
insuregeneticuniformity withineach strain.
Under the experimentalconditionsemployed,
these strainsretainedthe cannabinoid character
of theirseeds. Strain 152 (drug) could be dis-
tinguishedby a highlevel of A9-THC witha low
CBD level. Strain79 (non-drug) containedlow
levels of A9-THC but high levels of CBD, and
strain87 (fiber) also had low levels of A9-THC
and a high level of CBD. CBC and CBN were
foundin all threestrains,but in varyingamounts
while ?l8-THC was not detected. Similar results
were reportedby Small (1973) and others(Fet-
terman et al., 1971; Latta and Eaton, 1975)
which deemphasizesthe environmental influence
on total cannabinoidbiosynthesis.
3.8
In comparison,the distributionof the three
gland typesvariedfromclone to clone and subse-
quentlycould not be shownto fitany patternthat
mightcorrelatewith cannabinoid content. The
onlytrendobservedwas a decrease in both gland
numberand cannabinoidcontentas the plant or-
gan matured,but thiswas foundin all strainsand
in varyingdegrees. Comparisonof the analyses
of particularplant organs such as bracts and
leaves for each of these clones revealed a nega-
tive correlationbetweengland numberand can-
nabinoid content. Compared to the bract, floral
leaves subtendingthe bract have considerably
fewer glands per unit area, but only a slightly
lower cannabinoid content. Mature compound
leaves lacked capitate-stalkedglands as is the
case for most of the leaves on Cannabis except
the floralleaves subtendingthe bract. However,
comparisonof compoundleaves withfloralleaves
showedno outstandingdifferences thatmightim-
plicate capitate-stalkedglands as being solely re-
sponsiblefor cannabinoidcontent.
There are at least two explanationsforthe lack
of correlationbetweengland numberand canna-
binoid content. First, cannabinoids could pos-
sibly be present in other plant cells besides
glandular trichomes. A similar conclusion was
suggestedby Malingre et al. (1975) when he
found that leaf veins and pollen grains were
stainedby Fast Blue Salt B, a cannabinoidspecific
stain (DeFaubert Maunder, 1969).
Second, cannabinoidcontentof the individual
gland types could vary from one plant part to
anotherin responseto cannabinoidcharacter,en-
vironmentalconditions,gland aging,or any com-
bination of these. Differentamounts of canna-
TABLE 9. Cannabinoidcontentof capitatestalkedgland

ng Cannabinoid/gland
Gland
TABLE 7. Quantitationof glands on compoundleaves Clone age CBD A9-THC CBN

152 (drug) Mature * 57

Glands/mm2
Aged 35 21
Clone Capitate-stalkedCapitate-sessile Bulbous Senescent 9 1
152 (drug) 0 4.8 4.5 87 (fiber) Mature 229
Aged 113
79 (non-drug) 0 6.0 10.6 Senescent 29
87 (fiber) 0 2.6 8.1 *
Indicatesthe cannabinoidwas not detected.

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July,19771 TURNER ET AL -CANNABINOID CONTENT OF CANNABIS 693

binoids have been reportedto be presentin the CROMBIE, L., AND W. M. L. CROMBIE. 1975. Canna-
capitate-stalkedglands (Fairbairn, 1972; Malin- binoid formation in Cannabis sativa grafted inter-
gre et al., 1975). We found that mature,aged, racially, and with two Humulus species. Phyto-
and senescent glands had significantly chemistry
different DAYANANDAN, 14: 409-412.
P., AND P. B. KAUFMAN. 1976. Tri-
cannabinoid levels. However, at each stage of chomes of Cannabis sativa L. (Cannabaceae).
gland maturitythe cannabinoidcharacterof the Amer. J. Bot. 63: 578-591.
clone was maintained. Furthermore,ratios of DEFAUBERT MAUNDER, M. J. 1969. A simple and
mature,aged, and senescentglands were found specific test for Cannabis. J. Assoc. Public Anal.
to vary on each bract examined. It is possible 7: 24-30.
thatthe othergland typesalso may be subjectto DEPASQUALE, A. 1974. Ultrastructure of the Canna-
a similarvariabilityin theirconcentrationon the bis sativa glands. Planta Med. 25: 238-248.
DOORENBOS, N. J.,P. S. FETrERMAN, M. W. QUIMBY,
bract.
AND C. E. TURNER. 1971. Cultivation, extraction,
This investigation has demonstrateda negative and analysis of Cannabis sativa L. Annals N.Y.
correlation between cannabinoid content and Acad. Sci. 191: 3-14.
gland numberin a comparisonof different plant FAIRBA1N, J. W. 1972. The trichomes and glands of
organsfromthreephenotypically differentstrains Cannabissativa L. Bull. Narc. 24: 29-33.
maintainedas clones. Variation in numberand FETTERMAN, P. S., E. S. KEITH, C. W. WALLER, 0.
distributionof glands, as related to plant parts GUERRERO, N. J.DOORENBOS, AND M. W. QUIMBY.
and the age of those plant parts, emphasizes a 1971. Mississippi-grown Cannabis sativa L.: Pre-
need for cautionin employingthese structuresin liminary observation on chemical definition of
Cannabis systematics.These resultsalso empha- phenotype and variations in tetrahydrocannabinol
content versus age, sex, and plant part. J. Pharm.
size that stringentsamplingproceduresmust be
Sci. 60: 1246-1249.
employedas related to plant parts selected, age HAMMOND, C. T., AND P. G. MAHLBERG. 1973. Mor-
of plant parts, and presence and age of glands phology of glandular hairs of Cannabissativa from
on the plant parts,if valid comparativedata are scanning electron microscopy. Amer. J. Bot. 60:
being soughtfor cannabinoidsnot only between 524-528.
different strainsor withina singlestrain,but even HORRIDGE, G. A., AND S. L. TAMM. 1969. Critical
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