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CHINESE archaeological hemp?
- Thread starter Sam_Skunkman
- Start date
They used Cannabis, not smoked Cannabis, except for the Scythians and Pazyryk culture that smoked Cannabis by tossing it on coals under a tent, at least sometimes.
Archaeologist Sergei Ivanovich Rudenko unearthed tombs in the 1920's.
-SamS
Archaeologist Sergei Ivanovich Rudenko unearthed tombs in the 1920's.
-SamS
FOE20 - Here's some more info... I've posted this a few other places with zero reception. I guess people don't want to say "Cannabis Indica Indica" for a sativas and "Cannabis Indica Afghanica" for indicas.
This is from an interview w/ Robert Connell Clarke.
http://www.norml.org.nz/article509.html
**
This is from an interview w/ Robert Connell Clarke.
http://www.norml.org.nz/article509.html
**
Last edited by a moderator:
werd 3D_Dream Ive got that and the 2nd quote you show is from Hilligs report which has been publish already but yes great info..
I see it like theres aprox 5 types...Indica, Sativa mainly if were talking Palnst with Effects....Hemp as a 3rd but since it has no effect or terpinoid than is it a Cannabis or Hemp?....Then 2 more based on descendants of the main 2 varieties but if you take out the Hemp Factor....what do you get...Ind, Sats with the in between..but nothing that simple ever is it..heh
But instead of history spelling it out I'd like to distinguish on my own using the info these gentz have provided..
FOE20
I see it like theres aprox 5 types...Indica, Sativa mainly if were talking Palnst with Effects....Hemp as a 3rd but since it has no effect or terpinoid than is it a Cannabis or Hemp?....Then 2 more based on descendants of the main 2 varieties but if you take out the Hemp Factor....what do you get...Ind, Sats with the in between..but nothing that simple ever is it..heh
But instead of history spelling it out I'd like to distinguish on my own using the info these gentz have provided..
FOE20
M
mosca negra
Very interesting data. Always willing to listen to anything Rob has to say!
Genetic evidence for speciation in Cannabis (Cannabaceae)
karl Hillig
http://www.springerlink.com/content/t9367000120x1834/
karl Hillig
http://www.springerlink.com/content/t9367000120x1834/
Last edited:
B
bocil
Is an interesting report,many thanks Mr. Skunkman.
K
kopite
great read thx
lol i see you'd put the link in post 6 already foe...
I'd go with evolved by region, climate (enviroment), you either adapt/evole or you die... we all just want to survive
lol i see you'd put the link in post 6 already foe...
I'd go with evolved by region, climate (enviroment), you either adapt/evole or you die... we all just want to survive
Last edited:
G
Guest
Cool history here folks!
Cool history here folks!
Good work all,i admit i dont read many books..
But since you are talking asian weed,can i post some!
A bit of a break from the reading....my THAI plants going now.
Wish me luck.......they will go untill december friends!!!
Check the finish at the outdoor forum here at ic............
Cool history here folks!
Good work all,i admit i dont read many books..
But since you are talking asian weed,can i post some!
A bit of a break from the reading....my THAI plants going now.
Wish me luck.......they will go untill december friends!!!
Check the finish at the outdoor forum here at ic............
Last edited:
yo Pops..they took out 10 pgs..Its a 20pg report..I'll try n get it all up in here...
Sam thats the same link I used to use or actually there was a free one somewhere to...that one will cost $24 even for the short ver..I think...its all good tho..
hey Chimera n all I'll try n post this note for note...
nice LouGrew....look Thaialicious....hehe..
I\ll have to print screen a few images but I;ll add them later heres a clip that RevX had posted on another site that got my interest in the report..
I did highlight a few points I always consider to be quite to the point..
"considered to be only semi-domesticated"....for one..
FOE20
I have 2 ver..the 10pg and the 20pg..It seems the 20pg ver has a few more details and graphs they cut from the 10pg..
If you guys have a Email or a place I can upld or ftp a copy of the full PDF msg me or give me a link..no prob..Im just gana post the 20pg here or as much as possible..
----------------------------------------------------------------------------
Genetic Resources and Crop Evolution 52: 161–180, 2005. # Springer 2005
_____________________________________________________________
Genetic evidence for speciation in Cannabis (Cannabaceae)
Karl W. Hillig
Department of Biology, Indiana University, Bloomington, IN, USA; Current address: 1010 Saratoga Road,
Ballston Lake, NY 12019, USA (e-mail: [email protected])
Received 7 January 2003; accepted in revised form 28 June 2003
Key words: Allozyme, Cannabis, Evolution, Genetics, Origin, Taxonomy
Abstract
Sample populations of 157 Cannabis accessions of diverse geographic origin were surveyed for allozyme
variation at 17 gene loci. The frequencies of 52 alleles were subjected to principal components analysis. A
scatter plot revealed two major groups of accessions. The sativa gene pool includes fiber/seed landraces from
Europe, Asia Minor, and Central Asia, and ruderal populations from Eastern Europe. The indica gene pool
includes fiber/seed landraces from eastern Asia, narrow-leafleted drug strains from southern Asia, Africa,
and Latin America, wide-leafleted drug strains from Afghanistan and Pakistan, and feral populations from
India and Nepal. A third putative gene pool includes ruderal populations from Central Asia. None of the
previous taxonomic concepts that were tested adequately circumscribe the sativa and indica gene pools.
A polytypic concept of Cannabis is proposed, which recognizes three species, C. sativa, C. indica and
C. ruderalis, and seven putative taxa.
Abbreviations: PCA – principal components analysis
Introduction
Cannabis is believed to be one of humanity’s oldest cultivated crops, providing a source of fiber, food, oil, medicine, and inebriant since Neolithic times
(Chopra and Chopra 1957; Schultes 1973; Li 1974; Fleming and Clarke 1998). Cannabis is normally a dioecious, wind-pollinated, annual herb, although
plants may live for more than a year in subtropical regions (Cherniak 1982), and monoecious plants occur in some populations (Migal 1991).
The indigenous range of Cannabis is believed to be in Central Asia, the northwest Himalayas, and possibly extending into China (de Candolle 1885; Vavilov 1926;
Zhukovsky 1964; Li 1974). The genus may have two centers of diversity, Hindustani and European–Siberian (Zeven and Zhukovsky 1975).
Cannabis retains the ability to escape from cultivation and return to a weedy growth habit, and is considered to be only semi-domesticated (Vavilov 1926; Bredemann et al. 1956). Methods of Cannabis cultivation are described in the ancient literature
of China, where it has been utilized continuously for at least six thousand years (Li 1974). The genus may have been introduced into Europe ca. 1500
B.C. by nomadic tribes from Central Asia (Schultes 1970). Arab traders may have introduced Cannabis into Africa, perhaps one to two thousand
years ago (Du Toit 1980).
The genus is now distributed worldwide from the equator to about 60 N latitude, and throughout much of the southern hemisphere.
Cannabis cultivated for fiber and/or achenes (i.e., ‘seeds’) is herein referred to as ‘hemp.’ Cannabis breeders distinguish eastern Asian hemp from the
common hemp of Europe (Bo´csa and Karus 1998; de Meijer 1999). Russian botanists recognize four ‘eco-geographical’ groups of hemp: Northern,Middle-Russian, Southern, and Far Eastern (Serebriakova and Sizov 1940; Davidyan 1972). The Northern hemp landraces are smaller in stature
and earlier maturing than the landraces from more southerly latitudes, with a series of overlapping gradations in phenotypic traits between
theNorthern, Middle-Russian, and Southern types.
The Far-east Asian hemp landraces are most similar to the Southern eco-geographical group (Dewey1914). Two basic types of drug plant are commonly distinguished, in accord with the taxonomic
concepts of Schultes et al. (1974) and Anderson
(1980):
the narrow-leafleted drug strains and
the wide-leafleted drug strains
(Cherniak 1982; Anonymous 1989; de Meijer 1999).
The taxonomic treatment of Cannabis is problematic.
Linnaeus considered the genus to consist of a single undivided species, Cannabis sativa L.
Lamarck (1785) determined that Cannabis strains
from India are distinct from the common hemp
of Europe, and named the new species
C. indica Lam.
Distinguishing characteristics include more branching, a thinner cortex, narrower leaflets, and the general ability of C. indica to induce a state of
inebriation. Opinions differ whether Lamarck adequately differentiated
C. indica from
C. sativa, but they are both validly published species. Other species
of Cannabis have been proposed (reviewed in Schultes et al. 1974; and Small and Cronquist 1976), including C. chinensis Delile, and C. ruderalis
Janisch. Vavilov (1926) considered
C. ruderalis
to be synonymous with his own concept of C. sativa
L. var. spontanea Vav.
He later recognized wild Cannabis populations in....
Afghanistan to be distinct from
C. sativa var. spontanea,
and named the new taxon
C. indica Lam. var. kafiristanica Vav.
(Vavilov and Bukinich 1929).
Small and Cronquist (1976) proposed a monotypic treatment of Cannabis, which is a modification of the concepts of Lamarck and Vavilov. They
reduced C. indica in rank to C. sativa L. subsp. indica (Lam.) Small and Cronq. and differentiated it from C. sativa L. subsp. sativa, primarily on the
basis of ‘intoxicant ability’ and purpose of cultivation. Small and Cronquist bifurcated both subspecies into ‘wild’ (sensu lato) and domesticated
varieties on the basis of achene size, and other achene characteristics. This concept was challenged by other botanists, who used morphological traits
to delimit three species:
C. indica
C. sativa
C. ruderalis
(Anderson 1974, 1980; Emboden 1974;Schultes et al. 1974). Schultes et al. and Anderson
narrowly circumscribed C. indica to include relatively short, densely branched, wide-leafleted strains from Afghanistan.
The differences of opinion between taxonomists supporting monotypic
and polytypic concepts of Cannabis have not been
resolved (Emboden 1981).
Few studies of genetic variation in Cannabis have been reported. Lawi-Berger et al. (1982) studied seed protein variation in five fiber strains and
five drug strains of Cannabis, and found no basis for discriminating these predetermined groups.
de Meijer and Keizer (1996) conducted a more extensive investigation of protein variation in bulked seed lots of 147 Cannabis accessions, and on the basis of - five variable proteins concluded that
fiber
cultivars
fiber landraces
drug strains
and wild or naturalized populations could not be discriminated.
A method that shows greater promise for taxonomic investigation of Cannabis is random amplified polymorphic DNA (RAPD) analysis.
Using this technique, Cannabis strains from different geographic regions can be distinguished (Faetiet al. 1996; Jagadish et al. 1996; Siniscalco Gigliano
2001; Mandolino and Ranalli 2002),
but the number and diversity of accessions that have been analyzed
in these investigations are too small to provide a firm basis for drawing taxonomic inferences.
Allozyme analysis has proven useful in resolving difficult taxonomic issues in domesticated plants (Doebley 1989). Allozymes are enzyme variants
that have arisen through the process of DNA mutation. The genetic markers (allozymes) that are commonly assayed are part of a plant’s primary
metabolic pathways, and presumed neutral to the effects of human selection. Through allozyme analysis, it is possible to discern underlying patterns of
variation that have been outwardly obscured by the process of domestication. Because these genetic markers are cryptic, it is necessary to associate
allozyme frequencies with morphological differences in order to synthesize the genetic data into a formal taxonomic treatment (Pickersgill 1988).
Other types of biosystematic data may be included in the synthesis as well.
The purpose of this research is (1) to elucidate underlying genetic relationships among Cannabis accessions of known geographic origin, and (2) to assess previous taxonomic concepts in light of the genetic evidence. The research reported herein is part of a broader systematic investigation of morphological, chemotaxonomic, and genetic variation in Cannabis, which will be reported separately.
Materials and methods
The Cannabis germplasm collection
A diverse collection of 157 Cannabis accessions of
known geographic origin was obtained from breeders,
researchers, genebanks, and law enforcement
agencies (Table 1). Each accession consisted of an
unspecified number of viable achenes. Many of the
landraces that were studied are no longer cultivated,
and exist only in germplasm repositories.
Sixty-nine accessions were from hemp landraces
conserved at the N.I. Vavilov Institute of Plant
Industry (VIR) in Russia (Lemeshev et al. 1994).
Ten accessions were from Small’s taxonomic investigation
of Cannabis (Small and Beckstead 1973;
Small et al. 1976). Thirty-three accessions were
from de Meijer’s study of agronomic diversity in
Cannabis (de Meijer and van Soest 1992; de Meijer
1994, 1995; de Meijer and Keizer 1996). The
accessions from Afghanistan were obtained from
Cannabis breeders in Holland, and at least three
of these strains (Af-4, Af-5, Af-9) are inbred
(Anonymous 1989). Six Asian accessions were collected
from extant populations, including a drug
landrace from Pakistan (Pk-1), three feral populations
from India (In-2, In-3, In-5), and fiber
landraces from India (In-4) and China (Ch-4).
Accession Ch-4 was collected in Shandong
Province from seed propagated on the island of
Hunan (Clarke 1995). Five accessions from
Central Asia were collected from roadsides and
gardens in the Altai region of Russia, and identified
by the provider as C. ruderalis. Several
weedy accessions from Europe were identified as
C. ruderalis, ‘ssp. ruderalis,’ or ‘var. spontanea.’
A priori grouping of accessions
The accessions were assigned to drug or hempplantuse
groups, or ruderal (wild or naturalized) populations
as shown in Table 1. They were also assigned
to putative taxa according to the concepts of
Lamarck (1785), Delile (1849), Schultes et al.(1974) and Anderson (1980), and Small and
Cronquist (1976), based on morphological differences,
geographic origin, and presumed reason for
cultivation. Not all of the accessions could be unambiguously
assigned to a taxon for each concept. To
depict the various groups of interest, bivariate density
ellipses were drawn on the PC scatter plot. A
probability value of 0.75 was chosen because at this
value the ellipses encompass the majority of accessions
in a given group, but not the outliers.
Allozyme analysis
An initial survey was conducted to identify
enzymes that produce variable banding patterns
in Cannabis that can be visualized and interpreted
reliably(WendelandWeeden1989). Eleven enzymes
encoded at 17 putative loci were selected for a
genetic survey of the entire Cannabis germplasm
collection. Previously published methods of starch
gel electrophoresis and staining were employed
(Shields et al. 1983; Soltis et al. 1983; Morden et al.
1987; Wendel and Weeden 1989; Kephart 1990).
Gel/electrode buffer systems
Three gel/electrode buffer systems were utilized. A
Tris–citrate buffer system (modified from Wendel
and Weeden 1989) was used to resolve aconitase
(ACN), leucine aminopeptidase (LAP), malic
enzyme (ME), 6-phosphogluconate dehydrogenase
(6PGD), phosphoglucoisomerase (PGI), phosphoglucomutase
(PGM), and shikimate dehydrogenase
(SKDH). A lithium–borate buffer system
(modified from Soltis et al. 1983) was used to
resolve hexokinase (HK) and triosephosphate
isomerase (TPI). A morpholine–citrate buffer system
(modified fromWendel and Weeden 1989) was
used to resolve LAP, malate dehydrogenase
(MDH), ME, PGI, PGM, and an unknown
enzyme (UNK) that appeared on gels stained for
isocitrate dehydrogenase (IDH). IDH could not be
interpreted reliably, and was not used in the analysis.
A phosphate buffer (modified from Soltis et al.
1983) was used for enzyme extraction.
Electrophoresis and staining
For both the Tris–citrate and morpholine–citrate
buffer systems, 5-mm thick gels were held at 30 mA,
-------------------------------------------------------
and 10-mm thick gels at 45 mA throughout electrophoresis.
For the lithium–borate buffer system,
only 5-mm thick gels were used. These were held
at 50 mA for the first 10 min (after which the wicks
were removed), and at 200 V subsequently. Current
was applied for about 6 h to obtain good band
separation. Staining recipes for all enzymes except
HK were modified from Soltis et al. (1983). The
HK recipe was modified from Morden et al. (1987).
Tissue sample collection
Sample populations of each accession were grown
in two secure greenhouses at Indiana University,
Bloomington, Indiana. Voucher specimens are
deposited in the Deam Herbarium (IND) at
Indiana University. About 10 plants of each accession
were surveyed, except for accessions obtained
late in the investigation. Thirty Cannabis plants
were sampled for each gel. To make the gels easier
to interpret, two lanes were left blank or loaded
with a plant other than Cannabis. Tissue samples
were collected the afternoon before extraction and
electrophoresis, and stored overnight on moist filter
paper in small Petri dishes, under refrigeration.
Shoot tips generally produced the darkest bands,
although mature leaf tissue was better for visualizing
PGM.
Multivariate analysis
Putative genotypes were inferred from the allozyme
banding patterns, and allele frequencies were calculated
for small populations of each accession
(Wendel and Weeden 1989). Allele frequencies
were analyzed using JMP version 5.0 (SAS
Institute 2002). Principal components analysis
(PCA), commonly employed in numerical taxonomic
investigations, was used to visualize the
underlying pattern of genetic variation. The principal
components were extracted from the correlation
matrix of allele frequencies. Each PC axis is
defined by a linear combination of the allele frequencies.
PC axis 1 accounts for the largest amount
of variance that can be attributed to a single multivariate
axis, and each succeeding axis accounts for
a progressively smaller proportion of the remaining
variance. PC analysis simplifies the original
n-dimensional data set (n ¼ the number of alleles)
by enabling the data to be plotted on a reduced
number of orthogonal axes while minimizing the
loss of information. The degree of similarity among
the accessions can be inferred from their proximity
in PC space (Wiley 1981; Hillig and Iezzoni 1988).
The average number of alleles per locus (A),
number of alleles per polymorphic locus (Ap),
and percent polymorphic loci (P) were calculated
for each accession, and the expected heterozygosity
(He) averaged over all loci was calculated using the
mean allele frequencies of each sample population,
for the 11 enzymes that were assayed (Nei 1987;
Doebley 1989).
Several industrial hemp strains developed in
European breeding programs were genetically
characterized, but excluded from the statistical
analysis because of their possible hybrid origin
(de Meijer and van Soest 1992; de Meijer 1995).
For the purpose of this investigation, an accession
was considered hybrid if the parental strains
came from more than one country. Nine Chinese
accessions from the VIR collection were excluded
because of suspected hybridization during seed
regeneration. Only accessions analyzed in this
investigation are shown in Table 1.
Results
Gel interpretation
The allozyme banding patterns were interpreted as
shown in Figure 1. Only diploid banding patterns
were observed. When more than one set of
bands appeared on a gel, the loci were numbered
sequentially starting with the fastest migrating
(most anodal) locus. Alleles at a given locus were
lettered sequentially, starting with the fastest
migrating band. Monomeric enzymes (ACN, HK,
LAP, PGM, SKDH, UNK) showed a single band
for homozygous individuals, and two bands for
heterozygous individuals. Dimeric enzymes (6PGD,
MDH, PGI, TPI) typically showed one band for
homozygotes, and three bands for heterozygotes.
Malic enzyme (ME) is tetrameric (Weeden and
Wendel 1989), and heterozygous individuals produced
a five-banded pattern. Curiously, a pair of
bands appeared at the bottom of gels stained for
LAP due to cannabidiolic acid (CBDA) and tetrahydrocannabinolic
acid (THCA) migrating into
I'll add more tomorra.....
Sam thats the same link I used to use or actually there was a free one somewhere to...that one will cost $24 even for the short ver..I think...its all good tho..
hey Chimera n all I'll try n post this note for note...
nice LouGrew....look Thaialicious....hehe..
I\ll have to print screen a few images but I;ll add them later heres a clip that RevX had posted on another site that got my interest in the report..
I did highlight a few points I always consider to be quite to the point..
"considered to be only semi-domesticated"....for one..
FOE20
I have 2 ver..the 10pg and the 20pg..It seems the 20pg ver has a few more details and graphs they cut from the 10pg..
If you guys have a Email or a place I can upld or ftp a copy of the full PDF msg me or give me a link..no prob..Im just gana post the 20pg here or as much as possible..
----------------------------------------------------------------------------
Genetic Resources and Crop Evolution 52: 161–180, 2005. # Springer 2005
_____________________________________________________________
Genetic evidence for speciation in Cannabis (Cannabaceae)
Karl W. Hillig
Department of Biology, Indiana University, Bloomington, IN, USA; Current address: 1010 Saratoga Road,
Ballston Lake, NY 12019, USA (e-mail: [email protected])
Received 7 January 2003; accepted in revised form 28 June 2003
Key words: Allozyme, Cannabis, Evolution, Genetics, Origin, Taxonomy
Abstract
Sample populations of 157 Cannabis accessions of diverse geographic origin were surveyed for allozyme
variation at 17 gene loci. The frequencies of 52 alleles were subjected to principal components analysis. A
scatter plot revealed two major groups of accessions. The sativa gene pool includes fiber/seed landraces from
Europe, Asia Minor, and Central Asia, and ruderal populations from Eastern Europe. The indica gene pool
includes fiber/seed landraces from eastern Asia, narrow-leafleted drug strains from southern Asia, Africa,
and Latin America, wide-leafleted drug strains from Afghanistan and Pakistan, and feral populations from
India and Nepal. A third putative gene pool includes ruderal populations from Central Asia. None of the
previous taxonomic concepts that were tested adequately circumscribe the sativa and indica gene pools.
A polytypic concept of Cannabis is proposed, which recognizes three species, C. sativa, C. indica and
C. ruderalis, and seven putative taxa.
Abbreviations: PCA – principal components analysis
Introduction
Cannabis is believed to be one of humanity’s oldest cultivated crops, providing a source of fiber, food, oil, medicine, and inebriant since Neolithic times
(Chopra and Chopra 1957; Schultes 1973; Li 1974; Fleming and Clarke 1998). Cannabis is normally a dioecious, wind-pollinated, annual herb, although
plants may live for more than a year in subtropical regions (Cherniak 1982), and monoecious plants occur in some populations (Migal 1991).
The indigenous range of Cannabis is believed to be in Central Asia, the northwest Himalayas, and possibly extending into China (de Candolle 1885; Vavilov 1926;
Zhukovsky 1964; Li 1974). The genus may have two centers of diversity, Hindustani and European–Siberian (Zeven and Zhukovsky 1975).
Cannabis retains the ability to escape from cultivation and return to a weedy growth habit, and is considered to be only semi-domesticated (Vavilov 1926; Bredemann et al. 1956). Methods of Cannabis cultivation are described in the ancient literature
of China, where it has been utilized continuously for at least six thousand years (Li 1974). The genus may have been introduced into Europe ca. 1500
B.C. by nomadic tribes from Central Asia (Schultes 1970). Arab traders may have introduced Cannabis into Africa, perhaps one to two thousand
years ago (Du Toit 1980).
The genus is now distributed worldwide from the equator to about 60 N latitude, and throughout much of the southern hemisphere.
Cannabis cultivated for fiber and/or achenes (i.e., ‘seeds’) is herein referred to as ‘hemp.’ Cannabis breeders distinguish eastern Asian hemp from the
common hemp of Europe (Bo´csa and Karus 1998; de Meijer 1999). Russian botanists recognize four ‘eco-geographical’ groups of hemp: Northern,Middle-Russian, Southern, and Far Eastern (Serebriakova and Sizov 1940; Davidyan 1972). The Northern hemp landraces are smaller in stature
and earlier maturing than the landraces from more southerly latitudes, with a series of overlapping gradations in phenotypic traits between
theNorthern, Middle-Russian, and Southern types.
The Far-east Asian hemp landraces are most similar to the Southern eco-geographical group (Dewey1914). Two basic types of drug plant are commonly distinguished, in accord with the taxonomic
concepts of Schultes et al. (1974) and Anderson
(1980):
the narrow-leafleted drug strains and
the wide-leafleted drug strains
(Cherniak 1982; Anonymous 1989; de Meijer 1999).
The taxonomic treatment of Cannabis is problematic.
Linnaeus considered the genus to consist of a single undivided species, Cannabis sativa L.
Lamarck (1785) determined that Cannabis strains
from India are distinct from the common hemp
of Europe, and named the new species
C. indica Lam.
Distinguishing characteristics include more branching, a thinner cortex, narrower leaflets, and the general ability of C. indica to induce a state of
inebriation. Opinions differ whether Lamarck adequately differentiated
C. indica from
C. sativa, but they are both validly published species. Other species
of Cannabis have been proposed (reviewed in Schultes et al. 1974; and Small and Cronquist 1976), including C. chinensis Delile, and C. ruderalis
Janisch. Vavilov (1926) considered
C. ruderalis
to be synonymous with his own concept of C. sativa
L. var. spontanea Vav.
He later recognized wild Cannabis populations in....
Afghanistan to be distinct from
C. sativa var. spontanea,
and named the new taxon
C. indica Lam. var. kafiristanica Vav.
(Vavilov and Bukinich 1929).
Small and Cronquist (1976) proposed a monotypic treatment of Cannabis, which is a modification of the concepts of Lamarck and Vavilov. They
reduced C. indica in rank to C. sativa L. subsp. indica (Lam.) Small and Cronq. and differentiated it from C. sativa L. subsp. sativa, primarily on the
basis of ‘intoxicant ability’ and purpose of cultivation. Small and Cronquist bifurcated both subspecies into ‘wild’ (sensu lato) and domesticated
varieties on the basis of achene size, and other achene characteristics. This concept was challenged by other botanists, who used morphological traits
to delimit three species:
C. indica
C. sativa
C. ruderalis
(Anderson 1974, 1980; Emboden 1974;Schultes et al. 1974). Schultes et al. and Anderson
narrowly circumscribed C. indica to include relatively short, densely branched, wide-leafleted strains from Afghanistan.
The differences of opinion between taxonomists supporting monotypic
and polytypic concepts of Cannabis have not been
resolved (Emboden 1981).
Few studies of genetic variation in Cannabis have been reported. Lawi-Berger et al. (1982) studied seed protein variation in five fiber strains and
five drug strains of Cannabis, and found no basis for discriminating these predetermined groups.
de Meijer and Keizer (1996) conducted a more extensive investigation of protein variation in bulked seed lots of 147 Cannabis accessions, and on the basis of - five variable proteins concluded that
fiber
cultivars
fiber landraces
drug strains
and wild or naturalized populations could not be discriminated.
A method that shows greater promise for taxonomic investigation of Cannabis is random amplified polymorphic DNA (RAPD) analysis.
Using this technique, Cannabis strains from different geographic regions can be distinguished (Faetiet al. 1996; Jagadish et al. 1996; Siniscalco Gigliano
2001; Mandolino and Ranalli 2002),
but the number and diversity of accessions that have been analyzed
in these investigations are too small to provide a firm basis for drawing taxonomic inferences.
Allozyme analysis has proven useful in resolving difficult taxonomic issues in domesticated plants (Doebley 1989). Allozymes are enzyme variants
that have arisen through the process of DNA mutation. The genetic markers (allozymes) that are commonly assayed are part of a plant’s primary
metabolic pathways, and presumed neutral to the effects of human selection. Through allozyme analysis, it is possible to discern underlying patterns of
variation that have been outwardly obscured by the process of domestication. Because these genetic markers are cryptic, it is necessary to associate
allozyme frequencies with morphological differences in order to synthesize the genetic data into a formal taxonomic treatment (Pickersgill 1988).
Other types of biosystematic data may be included in the synthesis as well.
The purpose of this research is (1) to elucidate underlying genetic relationships among Cannabis accessions of known geographic origin, and (2) to assess previous taxonomic concepts in light of the genetic evidence. The research reported herein is part of a broader systematic investigation of morphological, chemotaxonomic, and genetic variation in Cannabis, which will be reported separately.
Materials and methods
The Cannabis germplasm collection
A diverse collection of 157 Cannabis accessions of
known geographic origin was obtained from breeders,
researchers, genebanks, and law enforcement
agencies (Table 1). Each accession consisted of an
unspecified number of viable achenes. Many of the
landraces that were studied are no longer cultivated,
and exist only in germplasm repositories.
Sixty-nine accessions were from hemp landraces
conserved at the N.I. Vavilov Institute of Plant
Industry (VIR) in Russia (Lemeshev et al. 1994).
Ten accessions were from Small’s taxonomic investigation
of Cannabis (Small and Beckstead 1973;
Small et al. 1976). Thirty-three accessions were
from de Meijer’s study of agronomic diversity in
Cannabis (de Meijer and van Soest 1992; de Meijer
1994, 1995; de Meijer and Keizer 1996). The
accessions from Afghanistan were obtained from
Cannabis breeders in Holland, and at least three
of these strains (Af-4, Af-5, Af-9) are inbred
(Anonymous 1989). Six Asian accessions were collected
from extant populations, including a drug
landrace from Pakistan (Pk-1), three feral populations
from India (In-2, In-3, In-5), and fiber
landraces from India (In-4) and China (Ch-4).
Accession Ch-4 was collected in Shandong
Province from seed propagated on the island of
Hunan (Clarke 1995). Five accessions from
Central Asia were collected from roadsides and
gardens in the Altai region of Russia, and identified
by the provider as C. ruderalis. Several
weedy accessions from Europe were identified as
C. ruderalis, ‘ssp. ruderalis,’ or ‘var. spontanea.’
A priori grouping of accessions
The accessions were assigned to drug or hempplantuse
groups, or ruderal (wild or naturalized) populations
as shown in Table 1. They were also assigned
to putative taxa according to the concepts of
Lamarck (1785), Delile (1849), Schultes et al.(1974) and Anderson (1980), and Small and
Cronquist (1976), based on morphological differences,
geographic origin, and presumed reason for
cultivation. Not all of the accessions could be unambiguously
assigned to a taxon for each concept. To
depict the various groups of interest, bivariate density
ellipses were drawn on the PC scatter plot. A
probability value of 0.75 was chosen because at this
value the ellipses encompass the majority of accessions
in a given group, but not the outliers.
Allozyme analysis
An initial survey was conducted to identify
enzymes that produce variable banding patterns
in Cannabis that can be visualized and interpreted
reliably(WendelandWeeden1989). Eleven enzymes
encoded at 17 putative loci were selected for a
genetic survey of the entire Cannabis germplasm
collection. Previously published methods of starch
gel electrophoresis and staining were employed
(Shields et al. 1983; Soltis et al. 1983; Morden et al.
1987; Wendel and Weeden 1989; Kephart 1990).
Gel/electrode buffer systems
Three gel/electrode buffer systems were utilized. A
Tris–citrate buffer system (modified from Wendel
and Weeden 1989) was used to resolve aconitase
(ACN), leucine aminopeptidase (LAP), malic
enzyme (ME), 6-phosphogluconate dehydrogenase
(6PGD), phosphoglucoisomerase (PGI), phosphoglucomutase
(PGM), and shikimate dehydrogenase
(SKDH). A lithium–borate buffer system
(modified from Soltis et al. 1983) was used to
resolve hexokinase (HK) and triosephosphate
isomerase (TPI). A morpholine–citrate buffer system
(modified fromWendel and Weeden 1989) was
used to resolve LAP, malate dehydrogenase
(MDH), ME, PGI, PGM, and an unknown
enzyme (UNK) that appeared on gels stained for
isocitrate dehydrogenase (IDH). IDH could not be
interpreted reliably, and was not used in the analysis.
A phosphate buffer (modified from Soltis et al.
1983) was used for enzyme extraction.
Electrophoresis and staining
For both the Tris–citrate and morpholine–citrate
buffer systems, 5-mm thick gels were held at 30 mA,
-------------------------------------------------------
and 10-mm thick gels at 45 mA throughout electrophoresis.
For the lithium–borate buffer system,
only 5-mm thick gels were used. These were held
at 50 mA for the first 10 min (after which the wicks
were removed), and at 200 V subsequently. Current
was applied for about 6 h to obtain good band
separation. Staining recipes for all enzymes except
HK were modified from Soltis et al. (1983). The
HK recipe was modified from Morden et al. (1987).
Tissue sample collection
Sample populations of each accession were grown
in two secure greenhouses at Indiana University,
Bloomington, Indiana. Voucher specimens are
deposited in the Deam Herbarium (IND) at
Indiana University. About 10 plants of each accession
were surveyed, except for accessions obtained
late in the investigation. Thirty Cannabis plants
were sampled for each gel. To make the gels easier
to interpret, two lanes were left blank or loaded
with a plant other than Cannabis. Tissue samples
were collected the afternoon before extraction and
electrophoresis, and stored overnight on moist filter
paper in small Petri dishes, under refrigeration.
Shoot tips generally produced the darkest bands,
although mature leaf tissue was better for visualizing
PGM.
Multivariate analysis
Putative genotypes were inferred from the allozyme
banding patterns, and allele frequencies were calculated
for small populations of each accession
(Wendel and Weeden 1989). Allele frequencies
were analyzed using JMP version 5.0 (SAS
Institute 2002). Principal components analysis
(PCA), commonly employed in numerical taxonomic
investigations, was used to visualize the
underlying pattern of genetic variation. The principal
components were extracted from the correlation
matrix of allele frequencies. Each PC axis is
defined by a linear combination of the allele frequencies.
PC axis 1 accounts for the largest amount
of variance that can be attributed to a single multivariate
axis, and each succeeding axis accounts for
a progressively smaller proportion of the remaining
variance. PC analysis simplifies the original
n-dimensional data set (n ¼ the number of alleles)
by enabling the data to be plotted on a reduced
number of orthogonal axes while minimizing the
loss of information. The degree of similarity among
the accessions can be inferred from their proximity
in PC space (Wiley 1981; Hillig and Iezzoni 1988).
The average number of alleles per locus (A),
number of alleles per polymorphic locus (Ap),
and percent polymorphic loci (P) were calculated
for each accession, and the expected heterozygosity
(He) averaged over all loci was calculated using the
mean allele frequencies of each sample population,
for the 11 enzymes that were assayed (Nei 1987;
Doebley 1989).
Several industrial hemp strains developed in
European breeding programs were genetically
characterized, but excluded from the statistical
analysis because of their possible hybrid origin
(de Meijer and van Soest 1992; de Meijer 1995).
For the purpose of this investigation, an accession
was considered hybrid if the parental strains
came from more than one country. Nine Chinese
accessions from the VIR collection were excluded
because of suspected hybridization during seed
regeneration. Only accessions analyzed in this
investigation are shown in Table 1.
Results
Gel interpretation
The allozyme banding patterns were interpreted as
shown in Figure 1. Only diploid banding patterns
were observed. When more than one set of
bands appeared on a gel, the loci were numbered
sequentially starting with the fastest migrating
(most anodal) locus. Alleles at a given locus were
lettered sequentially, starting with the fastest
migrating band. Monomeric enzymes (ACN, HK,
LAP, PGM, SKDH, UNK) showed a single band
for homozygous individuals, and two bands for
heterozygous individuals. Dimeric enzymes (6PGD,
MDH, PGI, TPI) typically showed one band for
homozygotes, and three bands for heterozygotes.
Malic enzyme (ME) is tetrameric (Weeden and
Wendel 1989), and heterozygous individuals produced
a five-banded pattern. Curiously, a pair of
bands appeared at the bottom of gels stained for
LAP due to cannabidiolic acid (CBDA) and tetrahydrocannabinolic
acid (THCA) migrating into
I'll add more tomorra.....
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hey fellaz....Tell me if this works...If so than grab at it...dl the shit out of it,.I did DL it 1x already to test and it looks good.....enjoy it..
FOE20
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http://www.4shared.com/file/61882707/dce30247/_3__genetic_evidence_for_speciation_in_cannabis.html
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FOE20
_____________________________________________________________________________________
http://www.4shared.com/file/61882707/dce30247/_3__genetic_evidence_for_speciation_in_cannabis.html
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I believe cannabis may have prompted our earliest ancestors to think deeply enough to begin making major advances in human history.
Moses' burning bush... well... hehehe He was a Shaman in Indica country. You KNOW he went up the mountain, to where the resins are best, and talked to the burning bush (the bush that you burn, NOT a bush that happens to be on fire and has conversations with folks as they walk up mountains).
I've often wondered if our earliest ancestors were given a bump in intelligence by discovering and consuming cannabis. The world's earliest civilizations ALL rose up where cannabis grew naturally.
Imagine an ape-like man, sitting in the shade of a tree with his family, eating cannabis... usually all he'd think is... eat... drink... sex... run... etc... But after eating cannabis flowers, he begins to conceive new thoughts... ways to make life easier, new ways to think about life.
I truly believe this may have been the case, or something like it.
Solomon, the wisest of kings, smoked cannabis. He loved it so much, he was buried with his stash! hehehe That's why so many reggae songs mention Solomon, or Solomon's grave.
Think about it... The VERY first humanoid to see a cannabis plant would have been intrigued by the shiny crystals, the amazing smells, and if he consumed it... He would never forget it, and try to find ways to keep a supply of it around.
Cannabis has been as much a part of the human experience as anything. It was probably the original "forbidden fruit", the fruit which Adam and Eve, the first thinking humans ate, which made them more aware of themselves than humans ever had been. I believe the Adam and Eve story may actually be the story of drug use, when the first humans realized there was some mystery to the world around them.
P.S. I don't believe the Adam and Eve story myself. I DO believe, however, that it's a story explaining how humans turned from simple, survival-minded beings, into something more.
Moses' burning bush... well... hehehe He was a Shaman in Indica country. You KNOW he went up the mountain, to where the resins are best, and talked to the burning bush (the bush that you burn, NOT a bush that happens to be on fire and has conversations with folks as they walk up mountains).
I've often wondered if our earliest ancestors were given a bump in intelligence by discovering and consuming cannabis. The world's earliest civilizations ALL rose up where cannabis grew naturally.
Imagine an ape-like man, sitting in the shade of a tree with his family, eating cannabis... usually all he'd think is... eat... drink... sex... run... etc... But after eating cannabis flowers, he begins to conceive new thoughts... ways to make life easier, new ways to think about life.
I truly believe this may have been the case, or something like it.
Solomon, the wisest of kings, smoked cannabis. He loved it so much, he was buried with his stash! hehehe That's why so many reggae songs mention Solomon, or Solomon's grave.
Think about it... The VERY first humanoid to see a cannabis plant would have been intrigued by the shiny crystals, the amazing smells, and if he consumed it... He would never forget it, and try to find ways to keep a supply of it around.
Cannabis has been as much a part of the human experience as anything. It was probably the original "forbidden fruit", the fruit which Adam and Eve, the first thinking humans ate, which made them more aware of themselves than humans ever had been. I believe the Adam and Eve story may actually be the story of drug use, when the first humans realized there was some mystery to the world around them.
P.S. I don't believe the Adam and Eve story myself. I DO believe, however, that it's a story explaining how humans turned from simple, survival-minded beings, into something more.
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Crazy Composer,
While I respect your right to believe what you like, I have seen no proof that Moses used Cannabis.
And as for Solomon, he smoked Cannabis? And was buried with his stash? Ok, where was Solomon buried?
And Adam and Eve were eating Cannabis?
I hate to shoot down your balloons but where is any proof at all of any of this?
I like the Space Bros story about them bringing Cannabis seeds to earth and giving it to Paleolithic man, this is pretty much a proven fact I thought....
I am not saying Cannabis did not effect human consciousness evolution, I am saying that there is zero proof that Cannabis was used by Moses, Solomon, Adam & Eve or any other Judeo-Christian personage in the Bible, or Tanakh, or new or old testament.
And people's desire to find one does not make it so.
-SamS
While I respect your right to believe what you like, I have seen no proof that Moses used Cannabis.
And as for Solomon, he smoked Cannabis? And was buried with his stash? Ok, where was Solomon buried?
And Adam and Eve were eating Cannabis?
I hate to shoot down your balloons but where is any proof at all of any of this?
I like the Space Bros story about them bringing Cannabis seeds to earth and giving it to Paleolithic man, this is pretty much a proven fact I thought....
I am not saying Cannabis did not effect human consciousness evolution, I am saying that there is zero proof that Cannabis was used by Moses, Solomon, Adam & Eve or any other Judeo-Christian personage in the Bible, or Tanakh, or new or old testament.
And people's desire to find one does not make it so.
-SamS
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Moses was an Egyptian shaman. He would have used cannabis as surely as he would have used other resins indigenous to the region, like frankincense and myhrr. If Moses was like other shaman of his time, he carried a special tent just for burning these resins and clambaking in the smoke... to commune with God. I can't believe you don't know this stuff.
Here's some interesting reading for you. I have read this kind of thing over and over and over for many years. I can't believe you haven't read any of this, Sam. Seems to me, someone like you ought to know as much as possible about this herb's historical uses. I guess we can all learn more than we currently know, eh?
http://cannabisculture.com/backissues/mayjune96/kanehbosm.html
^^^Look up Kaneh Bosm and you'll see for yourself that there's more to the story than meets your eye.
The word "Christ" itself means "the anointed one". Recipes have been found which show that as much as 9 pounds of cannabis were infused into oils and rubbed on the skin of Jesus AND Solomon to baptize them. Why do you think Jesus saw angels when John baptized him, and was tripping balls as John washed the oil from his skin.
Why do you think Solomon (when he was anointed king as a very young man) gained conciousness the morning after the anointing... under a cart in the market? It was good shit, that's why.
So, I have provided more proof for why it may have been so than you have provided for why it was not so. I await some equal evidence to the contrary.
Peace, good will,
cc
P.S. I believe the Adam and Eve story is a metaphor for the original humans becoming enlightened... To believe they were the very first humans is a leap of faith I'm not willing to stoop to. Many bible stories were written as metaphors, taken as literal tellings only by hardcore believers in modern times. People hearing the stories a thousand years ago would have known better than to take this stuff too literally.
Here's some interesting reading for you. I have read this kind of thing over and over and over for many years. I can't believe you haven't read any of this, Sam. Seems to me, someone like you ought to know as much as possible about this herb's historical uses. I guess we can all learn more than we currently know, eh?
http://cannabisculture.com/backissues/mayjune96/kanehbosm.html
^^^Look up Kaneh Bosm and you'll see for yourself that there's more to the story than meets your eye.
The word "Christ" itself means "the anointed one". Recipes have been found which show that as much as 9 pounds of cannabis were infused into oils and rubbed on the skin of Jesus AND Solomon to baptize them. Why do you think Jesus saw angels when John baptized him, and was tripping balls as John washed the oil from his skin.
Why do you think Solomon (when he was anointed king as a very young man) gained conciousness the morning after the anointing... under a cart in the market? It was good shit, that's why.
So, I have provided more proof for why it may have been so than you have provided for why it was not so. I await some equal evidence to the contrary.
Peace, good will,
cc
P.S. I believe the Adam and Eve story is a metaphor for the original humans becoming enlightened... To believe they were the very first humans is a leap of faith I'm not willing to stoop to. Many bible stories were written as metaphors, taken as literal tellings only by hardcore believers in modern times. People hearing the stories a thousand years ago would have known better than to take this stuff too literally.
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Here ya go, brother... So you don't have to look too deeply.
Of course there are scholars who deny it, they don't want the genesis of their (now cleaned up) religions to have anything to do with drugs. But look, the ill feelings toward hallucinogens and other drugs is a recent phenomena. Much respect has been (rightfully) given to mind opening/altering substances in ancient civilizations. The war on drugs is a new idea.
Did you know that Christmas is a shamanic, pagan, winter solstice holiday? In what is now called Russia, at the winter solstice, the celebration of the return of the Sun God, shaman would enter houses through holes in the roof (chimneys). And guess what they brought for everyone to celebrate with... Amanita Muscaria mushrooms. Men, women, children, grandparents, would consume the mushrooms and commune with the spirits. It was a jolly good time!
It's even thought by some that the three primary colors of Christmas, Green, red, and white, come from the color of the evergreen tree (under which these mushrooms grow), and the color of the mushrooms themselves (red and white). Red and white presents lain under the tree. Think about it. You never know... stranger truths exist all around us.
Did you know that Christmas is a shamanic, pagan, winter solstice holiday? In what is now called Russia, at the winter solstice, the celebration of the return of the Sun God, shaman would enter houses through holes in the roof (chimneys). And guess what they brought for everyone to celebrate with... Amanita Muscaria mushrooms. Men, women, children, grandparents, would consume the mushrooms and commune with the spirits. It was a jolly good time!
It's even thought by some that the three primary colors of Christmas, Green, red, and white, come from the color of the evergreen tree (under which these mushrooms grow), and the color of the mushrooms themselves (red and white). Red and white presents lain under the tree. Think about it.
You never know... stranger truths exist all around us.
