Cannabis genome
More actions
The cannabis genome is the full set of DNA carried by Cannabis sativa. Most of this DNA lies in the nucleus, in ten pairs of chromosomes. Small separate genomes lie in the mitochondria and chloroplasts. The plant is diploid (2n = 20), with nine pairs of autosomes and a pair of sex chromosomes, XX in females and XY in males.[1] One haploid set holds roughly 750 to 850 megabases (Mb), most of it repeated sequence.[2][3]
A draft sequence of the drug-type clone Purple Kush appeared in 2011. Long-read sequencing later gave chromosome-scale assemblies. By 2025 a pangenome joined 193 genomes from 144 plants.[4][3] Assembled length, gene count and repeat share differ from one assembly to the next. Most assemblies come from a narrow group of high-cannabinoid breeding lines. Asian material is underrepresented.[5][3] The wider study of heredity in the plant is the subject of cannabis genetics.
Chromosomes
The chromosomes are small, 2.6 to 3.8 micrometres long at metaphase. Most cannot be told apart by length and centromere position. In a 2014 karyotype of male and female hemp, eight autosome pairs were metacentric and one was submetacentric, the last bearing the nucleolus organiser with its 45S ribosomal genes. Another pair carried the 5S ribosomal genes. Fluorescent probes for the two sets of ribosomal genes and for CS-1, a repeat near the chromosome ends, picked out three of the nine autosome pairs. In relative length under the microscope, the Y came to 6.5 per cent, the X to 6.1 per cent and the largest autosome to 5.6 per cent. The 2014 karyotype classes both sex chromosomes as metacentric, where a 1998 study had called the X submetacentric and the Y subtelocentric.[1][2] Monoecious hemp cultivars have 2n = 20 with two X chromosomes and no Y. Their karyotypes vary widely within and between cultivars.[6]
Natural polyploids are rare. Of 483 plants from 84 accessions of wild or feral plants, landraces and cultivars, 482 were diploid. The one exception was a triploid male (2n = 30) from a wild North Indian accession. A tetraploid wild population from India had been reported once before. Other polyploids have been induced in breeding programmes.[7] Studies disagree over older duplications of the whole genome. Gao and colleagues inferred three such events from the JL assembly and Wei and colleagues two from the Yushe assembly. The authors of the pangenome found none since an ancient event about 100 million years ago and credited tandem and other local duplication for the plant's genomic diversity.[8][9][3]
Assemblies number their chromosomes differently, an inconsistency the authors of the Cannbio-2 assembly pointed out in 2020.[10] The cannabinoid synthase genes lie on chromosome 6 of the Purple Kush and Finola map and on chromosome 7 of the CBDRx assembly.[11][12] The sex chromosome pair is chromosome 1 in the 2018 version of CBDRx.[13] Later assemblies label the pair X and Y.[14][3]
Genome size
Flow cytometry in 1998, with Arabidopsis as the standard and DAPI as the stain, gave diploid genomes of 1,636 Mb for females and 1,683 Mb for males, a difference its authors thought might be due to the large Y.[2] Halved to 818 and 843 Mb, these are the values later assemblies quote as the expected size.[4][11][10]
A 2022 survey measured far more plants, from an area of more than 12,000 km across three continents. Diploids ranged from 1.810 to 2.152 picograms per diploid nucleus (pg/2C), a 1.189-fold spread with a mean of 1.956 pg/2C. Accession means differed by 1.073-fold. No accession or region could be clearly separated from the rest. The authors concluded that the plant's geographic spread and domestication had little effect on genome size. Earlier studies, averaging 1.720 pg/2C, had used other standards, instruments and stains, which the authors suggest may explain the gap. One of them found a 1.373-fold spread among only 35 plants.[7]
Males carry slightly more DNA than females: 1.987 against 1.947 pg/2C in the 2022 survey and 1.835 against 1.789 pg in a 2014 study, where monoecious plants matched females at 1.791 pg. Those 2014 values used soybean as the standard. With maize the same plants read 1.883, 1.835 and 1.838 pg. The authors suggested that the reference standard and the 1998 study's DAPI stain, which binds preferentially to AT bases, explain why their values ran higher than the 1998 ones. Male and female values overlap within accessions. The difference is too small for flow cytometry to separate the sexes when their nuclei are processed together.[7][15]
Sequence data give other figures. CBDRx alone has four. Counts of short DNA words (k-mers) in the reads predict 639 Mb. Its published assembly measures 737 Mb, of which 714 Mb is sequence anchored to chromosomes; with gaps counted, the same chromosomes come to 854 Mb.[12][10] The same kind of count gave 783 Mb for a female of the seed hemp Yushe and 816 Mb for a male.[9][16] The average haploid assembly in the pangenome is 781 Mb, while the same paper puts the genome at around 750 Mb.[3]
Assemblies
A genome assembly joins sequencing reads into longer stretches. Repeats and high heterozygosity make the job hard in cannabis.[8][3] Each row below gives what the study's own plant and methods produced. "Length" is the assembled length of one haploid set. Two values mean the two sets of one plant were assembled apart. Figures for repeats and genes were reached with different programs and are not strictly comparable.
| Published | Plant | Sequencing | Length (Mb) | Repeats (%) | Genes |
|---|---|---|---|---|---|
| 2011[4] | Purple Kush, a female clone of a drug-type strain | Illumina short reads, 454 mate pairs | 534, in scaffolds spanning more than 786 | 30,074 transcript clusters | |
| 2019[11][10] | Purple Kush (female) and the hemp Finola (male) | PacBio long reads, placed on a genetic map of their 99 offspring | 639 and 784 | about 73 | |
| 2020[8] | JL, a female from the Kyirong Gully in Tibet, described as wild | PacBio, Hi-C | 808 | 74.75 | 38,828 |
| 2020[10] | Cannbio-2, a female with balanced CBD and THC | PacBio, ordered against CBDRx | 914 as a draft, 904 ordered | 36,632 | |
| 2021[12] | CBDRx, a highly inbred high-CBD cultivar | Nanopore, Hi-C, genetic map of Skunk #1 × Carmen | 737 | 63 | 42,052 |
| 2024[9] | Yushe, a female of a local seed hemp from Shanxi | PacBio, Hi-C | 794 | 84.03 (transposable elements) | 31,091 |
| 2024[14] | Pink Pepper, a CBD-rich clone | PacBio, Illumina, nanopore | 770 | 77.13 | 30,459 |
| 2025[5] | Seven genotypes, two of them XY males | PacBio HiFi, Omni-C | mean 754 with an X, 782 with a Y | 23,182–28,296 | |
| 2025[3][17] | EH23, a hybrid of Early Resin Berry × HO40, within a set of 193 genomes | PacBio HiFi, Hi-C (EH23) | 739.94 (EH23a); mean 781 across the set | mean 68 (transposable elements, 78 assemblies) | 29,766 (EH23a); about 35,000 per genome |
| 2026[16] | Yushe, a male | PacBio HiFi, ultra-long nanopore | 770 and 804 | 74.53 and 76.33 | 32,519 and 31,465 |
| 2026[18] | F1 of the Colombian landrace Punto Rojo × the Colorado CBD clone Cherry Pie | Nanopore, reads sorted by parent | 794 and 774 | ||
| 2026[17] | A female from an accession described as wild | PacBio HiFi, ultra-long nanopore, Hi-C | 755.28 and 750.88 | 79.49 and 79.39 | 31,109 and 30,952 |
The 2011 draft, built from short reads, fell into 136,290 scaffolds (ordered runs of sequence) with half of its length in scaffolds of 16.2 kilobases or longer.[11] Its authors judged that it took in the vast majority of the genes and of the non-repetitive genome.[4] The 2019 map joined far longer stretches, using long reads and crossovers traced through a cross between Purple Kush and Finola. The 2021 CBDRx assembly added Hi-C contact data to nanopore reads. Even so, 32 per cent of the CBDRx genome lay in stretches that its genetic map could not order, nearly all in what the authors took to be centromere regions.[11][12] A near-complete pair published in 2026, the two haplotypes of one plant, has hundreds to thousands of telomere repeats at both ends of seven chromosomes and very few at one end of the other three.[17]
The two sets of chromosomes in one plant can differ a great deal, which is why recent assemblies separate them into haplotypes. In the 2026 near-complete pair, the haplotypes differ at 3.7 million single bases and 139 Mb of one has no aligned counterpart in the other. Across the pangenome, single-base heterozygosity ran from 1 to 2.5 per cent, while variable and unalignable regions together averaged 20.6 per cent of genome length.[17][3]
Only 23 per cent of pangenome genes occur in all 193 genomes. A further 55 per cent occur in 95 to 99 per cent of them. Most gene families are captured by 100 to 125 genomes, but the count of distinct short DNA words goes on rising as genomes are added.[3] A later index of 217 assemblies shows the same rise. A 66-haplotype graph published in 2026 points the same way: the two haplotypes of Yunma-7, an open-pollinated hemp from Yunnan, added substantial variation to it.[5][19] The choice of reference changes mapping results. One data set for the flowering locus Early1 gave numerous small peaks over 40 Mb on CBDRx and Pink Pepper but a peak of about 2 Mb when realigned to four new haplotypes, which also exposed assembly errors in the two older references.[5]
The sampling is uneven. Many assemblies are breeding lines of the Oregon CBD programme. These carry fewest unique sequence words, mostly under a million per haplotype. Asian hemp assemblies carry over ten million.[5] The one wild Tibetan assembly, JL, stood apart from every domesticated and feral line in the pangenome, which led the pangenome's authors to suggest that wild relatives survive in remote parts of Asia. They also wrote that the work ahead needs a careful separation of wild from feral plants.[3] The Punto Rojo haplotype, from what its sequencers call a Colombian landrace, was reached through a hybrid with a modern clone.[18]
Repeats and the layout of the chromosomes
The repeat share depends on plant and method. Clustering raw short reads from Purple Kush, Finola and USO-31, without any assembly, gave 64 to 65 per cent.[20] Assemblies run from 63 per cent masked in CBDRx to 84 per cent transposable elements in female Yushe. The introduction of the pangenome paper puts transposable elements at about 79 per cent of the genome; its measured average across 78 chromosome-scale assemblies is 68 per cent.[12][9][3]
Long terminal repeat retrotransposons are the largest class. They cover 58 to 60 per cent of the 2026 near-complete pair, in which Copia elements make up about 25 per cent and Gypsy elements about 17 per cent. By read clustering the two came to 14 and 14.8 per cent of the genome.[17][20] How recently these elements were active is disputed. The 2018 read-clustering study saw recent expansion in some families, yet took their relatively high ages overall to suggest little recent turnover. The pangenome paper reports some 4 million years of activity with a recent burst, many intact elements inserted within the last 100,000 years and elements still being transcribed. Recombination can delete most of an element and leave a solo long terminal repeat. Such solo copies are common in cannabis. The same authors link them to a genome far smaller than the 1,700 to 2,700 Mb of the sister genus Humulus.[20][3]
Tandem repeats, copies lying end to end, mark the chromosome ends and the centromeres. Every chromosome ends in the Arabidopsis-type telomere repeat, TTTAGGG.[1][17] Many arms carry a subtelomeric repeat just inside it: CS-1, about 375 bases long in the study that named it, or a repeat of 345 to 355 bases in CBDRx that matched 14 per cent of reads.[1][12] The centromere repeat differs from study to study. CBDRx has a repeat of 224 bases that matched 17 per cent of reads. The pangenome has a sparse array of 237 bases, usually near the centromere regions. The 2026 near-complete pair has two unrelated units of 125 and 112 bases that often occur joined.[12][3][17] In 2022 fluorescent probes placed a repeat called CS-237, of 237 bases with subunits of 111 and 126, on two chromosome pairs only: near the centromere on one and with the 45S ribosomal genes on the other.[21] Ribosomal DNA makes up about 2 per cent of the genome by read clustering. The 45S and 5S genes were missing from the 2019 assemblies.[20][11]
Genes and crossovers crowd towards the chromosome ends. The centres are rich in repeats and poor in genes. Crossovers there are rare. The authors of the 2019 map compared this layout with that of maize, barley and wheat and, to their knowledge, found it unusual outside such grain crops.[11] A later linkage map from the F2 progeny of two assembled genotypes shows the same bias towards the ends.[5] Large rearrangements also separate plants: an average of 86 inversions per genome in the pangenome, from 200 bases to 25 Mb, covering up to 7 per cent of a genome. One inversion of 19.5 Mb on chromosome 1 holds about 1,200 genes, among them the flowering-time gene PRR3.[3]
Cannabinoid synthase region
THCA synthase, CBDA synthase and CBCA synthase make the acid forms of the main cannabinoids. The first two largely set a plant's chemotype. The 2011 draft found THCA synthase in the Purple Kush transcriptome and CBDA synthase in its place in Finola, but it was too fragmented to say whether the two were alleles of one gene or separate, linked genes.[4][11]
In the 2019 map, the Purple Kush scaffold of about 250 kilobases that carries THCA synthase and the Finola scaffold of about 750 kilobases that carries CBDA synthase share almost nothing beyond long terminal repeat elements. The CBDA synthase scaffold lies inside a central stretch of about 39 Mb with no crossover among 99 offspring. One crossover set the THCA synthase scaffold apart at its edge. The two genes therefore sit at separate loci. Purple Kush also held a CBDA synthase pseudogene interrupted by a Gypsy element. Finola held no THCA synthase at all. A THCA-synthase-like gene present in both proved to be CBCA synthase.[11]
The 2021 CBDRx assembly placed 13 synthase sequences on chromosome 7 near 26, 29 and 31 Mb, in a pericentromeric region of suppressed recombination: an array of seven, an array of five and a lone copy of CBDA synthase. Each copy sits in a repeating unit of 31 to 45 kilobases among long terminal repeat elements. Only the lone copy was highly expressed. Two of the three contigs bearing functional genes could not be ordered from the map and were placed side by side as the simplest arrangement, which single contigs in two related First Light plants later supported. By read depth the synthase copies number 25 in Purple Kush and 17 in Finola, of which 13 and 16 were assembled.[12] Cannbio-2 has 16 candidate genes, 12 of them on chromosome 7.[10]
Across the pangenome each haploid genome holds at most one full-length THCA or CBDA synthase, set among partial copies in a limited number of arrangements and confined to about 1.5 Mb of chromosome 7. Most of these full-length genes do not line up between cannabis types and are tied to inversions. CBCA synthase usually lies 15 to 20 Mb from the centromere, occurs in 56 per cent of genomes, in arrays of 1 to 15 copies, and is little expressed.[3] A comparison with related genera traces the family to duplications within the Cannabis lineage, out of a larger group of berberine-bridge-enzyme-like genes.[22]
The link between copy number and cannabinoid content is not settled. Genome data from 69 plants showed that synthase copy number varies, and the authors concluded that it partly explains content. In the 35 plants of 22 cultivars with matching chemical data, most correlations lost significance after correction for relatedness, while copy numbers of the THCA and CBDA synthase family stayed positively correlated with THC.[23] In the Skunk #1 × Carmen cross, none of six quantitative trait loci for total cannabinoids lay at the arrays, though one signal there fell just short of significance.[12] Close linkage of the two main genes accounts for the single-locus inheritance seen in crosses, a pattern described under genetic traits.[12] Complete THCA synthase is largely missing from hemp cultivars and complete CBDA synthase from drug cultivars. Plants of a group called basal in a 2021 resequencing study may carry either or both. Its authors read the pattern as artificial selection acting on an ancestor that had both genes.[24]
Sex chromosomes
The 2019 map could not place its male-specific markers. CBDRx and the other assemblies from female plants have no Y.[11][12] In 2020, sex-linked genes were identified from the transcripts of two parents and ten offspring. Most of those that could be placed mapped to chromosome 1 of the 2018 CBDRx assembly. That chromosome held an X-specific region from 30 to 105 Mb and a pseudoautosomal region from 1 to 30 Mb.[13] The first assembled Y chromosomes became public in 2024.[5]
In the pangenome the Y spans 108 to 113 Mb and the X 83 to 85 Mb. A pseudoautosomal region of about 29 Mb is followed by a sex-determining region of 79 to 84 Mb on the Y, with 840 to 1,160 genes, or an X-specific region of 53 to 55 Mb.[3] That X-specific region is shorter than the roughly 75 Mb inferred from the 2018 assembly. A male of Yushe assembled separately has an 84 Mb sex-determining region, 71 per cent of its Y, of which 91.4 per cent is repeats. It holds 570 protein-coding genes against 1,529 in the matching part of the X. It also has two inverted segments of 25.6 and 54.0 Mb.[16] The assembled Y is about 30 per cent longer than the X, while under the microscope the two differ far less, at 6.5 and 6.1 per cent of relative length.[3][1] Whole haploid sets differ by little: in two males sequenced in 2025, the set holding the Y was 3.93 and 6.02 per cent longer than the set holding the X.[5]
Estimates of the age of the sex chromosomes have risen. The 2020 study gave about 12 to 29 million years, from divergence between X and Y gene copies and two molecular clocks. Work on hop in 2021 showed the pair is shared with Humulus and put the start of suppressed recombination at 21 to 25 million years or more. A 2026 study of assembled X and Y chromosomes places their origin before the split of Cannabis and Humulus, around 36 million years ago, and gets 38.6 to 48.3 million years from gene pairs in hop. The Yushe male gave about 37.7 million years (31.1 to 44.3), calibrated on the split from mulberry.[13][25][26][16]
Which gene determines sex remains open. The pangenome paper leaves the male-determining genes of the Y unidentified. The Yushe study proposes a male-expressed gene on the Y. The 2026 study finds no clear sex-determining gene on the Y and proposes an X-linked ACS gene of the ethylene pathway as the candidate.[3][16][26]
Organelle genomes
The mitochondrial and chloroplast genomes are far smaller than the nuclear one. Sequencing of the hemp cultivar Carmagnola gave a mitochondrial genome of 415,499 base pairs. A later nanopore study gave 415,837 for its main form and 153,927 for the chloroplast genome.[27][28] Their inheritance and their use in studies of descent are covered under genetic marker.
See also
References
- ↑ 1.0 1.1 1.2 1.3 1.4 Divashuk, M.G.; Alexandrov, O.S.; Razumova, O.V.; Kirov, I.V.; Karlov, G.I. (2014). "Molecular cytogenetic characterization of the dioecious Cannabis sativa with an XY chromosome sex determination system". PLoS ONE. 9 (1) e85118. doi:10.1371/journal.pone.0085118.
- ↑ 2.0 2.1 2.2 Sakamoto, K.; Akiyama, Y.; Fukui, K.; Kamada, H.; Satoh, S. (1998). "Characterization; genome sizes and morphology of sex chromosomes in hemp (Cannabis sativa L.)". Cytologia. 63 (4): 459–464. doi:10.1508/cytologia.63.459.
- ↑ 3.00 3.01 3.02 3.03 3.04 3.05 3.06 3.07 3.08 3.09 3.10 3.11 3.12 3.13 3.14 3.15 3.16 3.17 3.18 Lynch, R.C.; Padgitt-Cobb, L.K.; Garfinkel, A.R.; Knaus, B.J.; Hartwick, N.T.; et al. (2025). "Domesticated cannabinoid synthases amid a wild mosaic cannabis pangenome". Nature. 643 (8073): 1001–1010. doi:10.1038/s41586-025-09065-0. PMC 12286863.
- ↑ 4.0 4.1 4.2 4.3 4.4 van Bakel, H.; Stout, J.M.; Cote, A.G.; Tallon, C.M.; Sharpe, A.G.; Hughes, T.R.; Page, J.E. (2011). "The draft genome and transcriptome of Cannabis sativa". Genome Biology. 12 (10) R102. doi:10.1186/gb-2011-12-10-r102.
- ↑ 5.0 5.1 5.2 5.3 5.4 5.5 5.6 5.7 Stack, G.M.; Quade, M.A.; Wilkerson, D.G.; Monserrate, L.A.; Bentz, P.C.; et al. (2025). "Comparison of recombination rate, reference bias, and unique pangenomic haplotypes in Cannabis sativa using seven de novo genome assemblies". International Journal of Molecular Sciences. 26 (3) 1165. doi:10.3390/ijms26031165. PMC 11818205.
- ↑ Razumova, O.V.; Alexandrov, O.S.; Divashuk, M.G.; Sukhorada, T.I.; Karlov, G.I. (2016). "Molecular cytogenetic analysis of monoecious hemp (Cannabis sativa L.) cultivars reveals its karyotype variations and sex chromosomes constitution". Protoplasma. 253 (3): 895–901. doi:10.1007/s00709-015-0851-0.
- ↑ 7.0 7.1 7.2 Balant, M.; Rodríguez González, R.; Garcia, S.; Garnatje, T.; Pellicer, J.; Vallès, J.; Vitales, D.; Hidalgo, O. (2022). "Novel insights into the nature of intraspecific genome size diversity in Cannabis sativa L." Plants. 11 (20) 2736. doi:10.3390/plants11202736.
- ↑ 8.0 8.1 8.2 Gao, Shan; Wang, Baishi; Xie, Shanshan; Xu, Xiaoyu; Zhang, Jin; Pei, Li; Yu, Yongyi; Yang, Weifei; Zhang, Ying (2020). "A high-quality reference genome of wild Cannabis sativa". Horticulture Research. 7 (1) 73. doi:10.1038/s41438-020-0295-3.
- ↑ 9.0 9.1 9.2 9.3 Wei, H.; Yang, Z.; Niyitanga, S.; Tao, A.; Xu, J.; Fang, P.; et al. (2024). "The reference genome of seed hemp (Cannabis sativa) provides new insights into fatty acid and vitamin E synthesis". Plant Communications. 5 (1) 100718. doi:10.1016/j.xplc.2023.100718.
- ↑ 10.0 10.1 10.2 10.3 10.4 10.5 Braich, S.; Baillie, R.C.; Spangenberg, G.C.; Cogan, N.O.I. (2020). "A new and improved genome sequence of Cannabis sativa". GigaByte. 2020: 1–13. doi:10.46471/gigabyte.10.
- ↑ 11.00 11.01 11.02 11.03 11.04 11.05 11.06 11.07 11.08 11.09 Laverty, K.U.; Stout, J.M.; Sullivan, M.J.; Shah, H.; Gill, N.; Holbrook, L.; Deikus, G.; Sebra, R.; Hughes, T.R.; Page, J.E.; van Bakel, H. (2019). "A physical and genetic map of Cannabis sativa identifies extensive rearrangements at the THC/CBD acid synthase loci". Genome Research. 29 (1): 146–156. doi:10.1101/gr.242594.118. PMC 6314170.
- ↑ 12.00 12.01 12.02 12.03 12.04 12.05 12.06 12.07 12.08 12.09 12.10 Grassa, C.J.; Weiblen, G.D.; Wenger, J.P.; Dabney, C.; et al. (2021). "A new Cannabis genome assembly associates elevated cannabidiol (CBD) with hemp introgressed into marijuana". New Phytologist. 230 (4): 1665–1679. doi:10.1111/nph.17243. PMC 8248131.
- ↑ 13.0 13.1 13.2 Prentout, D.; Razumova, O.; Rhoné, B.; Badouin, H.; Henri, H.; Feng, C.; Käfer, J.; Karlov, G.; Marais, G.A.B. (2020). "An efficient RNA-seq-based segregation analysis identifies the sex chromosomes of Cannabis sativa". Genome Research. 30 (2): 164–172. doi:10.1101/gr.251207.119.
- ↑ 14.0 14.1 Ryu, B.R.; Gim, G.J.; Shin, Y.R.; Kang, M.J.; Kim, M.J.; Kwon, T.H.; et al. (2024). "Chromosome-level haploid assembly of Cannabis sativa L. cv. Pink Pepper". Scientific Data. 11 (1) 1442. doi:10.1038/s41597-024-04288-8.
- ↑ Faux, A.-M.; Berhin, A.; Dauguet, N.; Bertin, P. (2014). "Sex chromosomes and quantitative sex expression in monoecious hemp (Cannabis sativa L.)". Euphytica. 196 (2): 183–197. doi:10.1007/s10681-013-1023-y.
- ↑ 16.0 16.1 16.2 16.3 16.4 Wei, H.; Yang, Z.; Zhuang, L.; Pan, X.; Jia, H.; Jiang, S.; et al. (2026). "A haplotype-resolved genome assembly of seed hemp (Cannabis sativa) and analysis of Y chromosome divergence from the X". Horticulture Research. 13 (1) uhaf268. doi:10.1093/hr/uhaf268.
- ↑ 17.0 17.1 17.2 17.3 17.4 17.5 17.6 Ni, Y.; Zhang, C.; Li, J.; Wu, W.; Shen, G.; Xie, Y.; Song, C.; Liu, C. (2026). "A near-complete, haplotype-resolved telomere-to-telomere genome assembly of Cannabis sativa reveals complex higher-order repetitive structures". Plant Communications. 7 (9) 101913. doi:10.1016/j.xplc.2026.101913.
- ↑ 18.0 18.1 Pike, B.; Kozik, A.; Terán, W. (2026). "A trio-binning approach for genome assembly reveals extensive structural variation between two Cannabis cultivars: Punto Rojo and Cherry Pie". G3: Genes, Genomes, Genetics. 16 (2) jkaf286. doi:10.1093/g3journal/jkaf286.
- ↑ Pike, B.; Gonçalves da Silva, A.; Terán, W. (2026). "Four haplotype-resolved genome assemblies and a reference-free 66-haplotype pangenome graph for Cannabis sativa". Scientific Data. 13 (1) 1232. doi:10.1038/s41597-026-07806-y.
- ↑ 20.0 20.1 20.2 20.3 Pisupati, R.; Vergara, D.; Kane, N.C. (2018). "Diversity and evolution of the repetitive genomic content in Cannabis sativa". BMC Genomics. 19 (1) 156. doi:10.1186/s12864-018-4494-3.
- ↑ Alexandrov, O.S.; Romanov, D.V.; Divashuk, M.G.; Razumova, O.V.; Ulyanov, D.S.; Karlov, G.I. (2022). "Study and physical mapping of the species-specific tandem repeat CS-237 linked with 45S ribosomal DNA intergenic spacer in Cannabis sativa L." Plants. 11 (11) 1396. doi:10.3390/plants11111396.
- ↑ van Velzen, R.; Schranz, M.E. (2021). "Origin and evolution of the cannabinoid oxidocyclase gene family". Genome Biology and Evolution. 13 (8) evab130. doi:10.1093/gbe/evab130.
- ↑ Vergara, D.; Huscher, E.L.; Keepers, K.G.; Givens, R.M.; Cizek, C.G.; Torres, A.; Gaudino, R.; Kane, N.C. (2019). "Gene copy number is associated with phytochemistry in Cannabis sativa". AoB PLANTS. 11 (6) plz074. doi:10.1093/aobpla/plz074.
- ↑ Ren, G.; Zhang, X.; Li, Y.; Ridout, K.; Serrano-Serrano, M.L.; Yang, Y.; Liu, A.; Ravikanth, G.; Nawaz, M.A.; Mumtaz, A.S.; Salamin, N.; Fumagalli, L. (2021). "Large-scale whole-genome resequencing unravels the domestication history of Cannabis sativa". Science Advances. 7 (29) eabg2286. doi:10.1126/sciadv.abg2286.
- ↑ Prentout, D.; Stajner, N.; Cerenak, A.; Tricou, T.; Brochier‐Armanet, C.; Jakse, J.; Käfer, J.; Marais, G.A.B. (2021). "Plant genera Cannabis and Humulus share the same pair of well-differentiated sex chromosomes". New Phytologist. 231 (4): 1599–1611. doi:10.1111/nph.17456.
- ↑ 26.0 26.1 Carey, S.B.; Bentz, P.C.; Lovell, J.T.; Akozbek, L.M.; Myers, Z.A.; Korani, W.; et al. (2026). "An X-linked sex determination mechanism in cannabis and hop". Nature Communications. 17 (1) 6866. doi:10.1038/s41467-026-73233-7.
- ↑ White, K.H.; Vergara, D.; Keepers, K.G.; Kane, N.C. (2016). "The complete mitochondrial genome for Cannabis sativa". Mitochondrial DNA Part B. 1 (1): 715–716. doi:10.1080/23802359.2016.1155083.
- ↑ Liu, J.; Ni, Y.; Liu, C. (2023). "Polymeric structure of the Cannabis sativa L. mitochondrial genome identified with an assembly graph model". Gene. 853 147081. doi:10.1016/j.gene.2022.147081.