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Official websites use. Share sensitive information only on official, secure websites. Corresponding authors: E-mails: zcxu nefu. Is Cannabis a boon or bane? Cannabis sativa has long been a versatile crop for fiber extraction industrial hemp , traditional Chinese medicine hemp seeds , and recreational drugs marijuana. A comprehensive understanding of the underlying mechanism of cannabinoid biosynthesis is necessary to cultivate and promote globally the medicinal application of Cannabis resources. Here, we comprehensively review the historical usage of Cannabis , biosynthesis of trichome-specific cannabinoids, regulatory network of trichome development, and synthetic biology of cannabinoids. This review provides valuable insights into the efficient biosynthesis and green production of cannabinoids, and the development and utilization of novel Cannabis varieties. Historically, Cannabis has been grown as an important food, fiber, and medicinal crop. The whole Cannabis plant is used for various purposes. For example, its stems have been used for fiber production, which has been practiced for millennia. Additionally, Cannabis roots are used to treat inflammation and pain \[ 4 \]. Cannabinoids, abundant in secretory glandular trichomes GTs densely inlaid in the female inflorescence of Cannabis \[ 5 \], exhibit medicinal or recreational properties. Cannabinoids, a group of C21 terpenophenolic compounds, are the most abundant in Cannabis. Further, cannabidiol CBD , a nonpsychoactive cannabinoid, has been clinically validated to treat specific medical conditions, such as epilepsy, glaucoma, and depressive disorder. However, the legal status and regulations surrounding the use of Cannabis and its cannabinoids vary by different countries. Therefore, it is essential to seek guidance from medical professionals and adhere to local laws and regulations when considering the medical applications of Cannabis. This growth also presents significant challenges for Cannabis cultivation and cannabinoid production. Remarkable advancements in the cultivation of Cannabis plants with high-yielding cannabinoids and the reconstruction of cannabinoid production in microorganisms via metabolic engineering have been achieved \[ 9 \]. The integration of multi-omics methodologies, including genomics, transcriptomics, and metabolomics, has provided comprehensive insights into the genetic composition, gene expression patterns, and regulation of cannabinoid biosynthesis. Further, the relaxation of regulations and the authorization of Cannabis research have expanded the opportunities for studying this plant. Here, we summarize Cannabis historical usage, cannabinoid biosynthesis, trichome development regulatory network, and cannabinoid metabolic engineering. This review aims to enhance the understanding of Cannabis development and utilization, paving the way for further exploration and innovation in Cannabis research. Cannabis has a long history of human use, with documented evidence from around the world \[ 10 , 11 \] Fig. The description of Soma in Rig Veda BCE , an ancient Indian literature, may be the earliest reference to the psychoactive activity of Cannabis \[ 13 \]. A series of recent events on the medicinal properties of Cannabis have gradually unraveled its pharmacological mechanism. In , Napoleon brought Cannabis back to France from Egypt and investigated it for its pain-relieving and sedative qualities \[ 15 \]. Subsequently, the discovery of CB1 and CB2 receptors, the target of cannabinoids in the human body, and endocannabinoid system ECS has revealed the relationship between cannabinoids and human health in maintaining homeostasis and influencing various functions such as sleep, appetite, pain perception, inflammation, memory, mood, and reproduction \[ 19 , 20 \]. Originally, Cannabis was primarily cultivated to obtain seeds and fiber \[ 21 \]. These ancient written records highlight the significance of hemp as an important textile material at that time. Additionally, increasing archaeological evidence indicated the spiritual use of Cannabis smoke before BCE. It is believed to be the earliest discovery of the narcotic and hallucinogenic effects of Cannabis \[ 22 \]. Cannabinol CBN , an oxidation product of THC, was detected on the surface of ten wooden fire pots and cobblestones excavated from the Jirzankal cemetery in the Pamir plateau BCE , suggesting the clear evidence of the burning of Cannabis \[ 23 \]. The psychoactive properties of marijuana have led to restrictions on its usage in various countries. In recent times, however, some countries and regions have begun liberalizing the restrictions on Cannabis for its economic and pharmacological benefits. California was the first US state to legalize the drug marijuana, ushering in the era of store-bought, commercially made edibles in \[ 24 \]. Colorado was the first state to legalize recreational marijuana, imposing a limit of 10 mg THC per serving for edibles in \[ 25 \]. In the same year, Uruguay became the first country to legalize Cannabis. Subsequently, Thailand was the first Asian country to legalize Cannabis in \[ 26 \]. The legalization has allowed the development of more diverse properties of Cannabis for industrial and medical usage. The origin and domestication of Cannabis are debated. Central and Southeast Asia have been proposed as potential regions for its natural origin and primary domestication, playing a significant role in its evolutionary history \[ 30 \]. However, recent microfossil fossil pollen data suggest a center of origin in the northeastern Tibetan Plateau \[ 7 \]. Further, studies examining the correlation between genetic and geographic distances based on the chloroplast genome propose that Cannabis likely originated in low-latitude regions \[ 31 \]. The complex genetic makeup of Cannabis , characterized by high heterozygosity, has posed challenges in studying its domestication history \[ 32 \]. According to achene fossils found in East Asia and Europe, early humans employed hemp as a fiber plant, whereas the ancient use of the drug marijuana dates back to at least years ago in Central Asia, as discovered via wooden pots containing THC, which were probably employed for ritualistic and medicinal purposes \[ 23 , 33 , 34 \]. However, large-scale whole-genome resequencing studies involving Cannabis resources from around the world suggest that Cannabis was domesticated in East Asia during the early Neogene \[ 35 \]. Over millennia, both artificial domestication and wild cultivation led to the development of diverse Cannabis species, which are cultivated for their fiber and as drugs Fig. S1, see online supplementary material. Cannabis contains numerous natural compounds \[ 36 \] and over compounds have been identified, such as cannabinoids, phenolics, terpenes, and alkaloids. Cannabinoids form the majority of compounds found in Cannabis , with more than cannabinoids having been isolated. Cannabinoids exist in two chemical forms: decarboxylated forms and carboxylated forms. In fresh Cannabis tissues, carboxylated form is the predominant form. However, through nonenzymatic reactions such as drying, aging, heating, or incineration, Cannabis and its extracts undergo decarboxylation of carboxylated forms to form decarboxylated forms \[ 39 \]. Cannabinoids and human health. A Eleven subclasses of cannabinoids in Cannabis and their respective pharmacological activities. THC and CBD are the most significant compounds in Cannabis , and their effects can be perceived as both positive and negative \[ 40 \]. Thus, some individuals may be more prone to the effects of THC, although it also exhibits antiepileptic, antitumor, and antiemetic effects \[ 43 , 44 \]. In contrast, CBD, a nonpsychoactive component of Cannabis , acts as an antagonist to the association between THC and the receptors, effectively mitigating the hallucinogenic effects of THC on the body. CBD can be used in the restoration of the ECS, helping alleviate physical ailments and emotional distress \[ 45 , 46 \] Fig. It has verified that CBD possesses diverse pharmacological effects, including antibacterial, anti-inflammatory, anxiolytic, and antiepileptic properties, and it is considered safe for humans \[ 47—49 \]. Recent studies have revealed promising findings regarding the therapeutic properties of various cannabinoids \[ 6 , 50 \]. For instance, CBC is reported to exert anti-inflammatory, antitumor, antidepressant, and antifungal effects \[ 51—54 \]. CBG shows potential as an antibiotic and has also been associated with antitumor, antidepressant, analgesic, and glaucoma-alleviating properties \[ 51 , 52 , 55 , 56 \]. Meanwhile, CBN stimulates appetite and possesses anti-asthma, tranquilizing, and pain-relieving properties \[ 51 , 52 , 57 \]. However, some of the lesser-known cannabinoids may be scarce and difficult to extract, resulting in limited research on their medicinal functions and functional activities. Further exploration is needed to fully understand their potential benefits. Cannabinoids are synthesized and accumulated in the Cannabis secretory GTs, and the content of cannabinoid is related to the secretion, type, and density of GTs \[ 5 \]. Until now, the skeleton biosynthesis of cannabinoids has been elucidated, including two main biosynthetic pathways Fig. The first pathway is polyketide synthesis occurring within the cytosol. It involves the conversion of hexanoic acid to thiolate hexanoyl coenzyme A by acyl-activating enzyme AAE \[ 58 \]. Olivetol synthase OLS then catalyzes hexanoyl coenzyme A condensation with malonyl coenzyme A, leading to the formation of an intermediate tetraketide-CoA \[ 59 \]. This intermediate is further cyclized by olivetolic acid cyclase OAC to produce olivetolic acid OA , which serves as the polyketide nucleation component required for cannabinoid synthesis \[ 60 , 61 \]. The second pathway is the methylerythritol 4-phosphate MEP pathway, which generates isopentenyl diphosphate and dimethyl allyl diphosphate. These two compounds combine to form geranyl pyrophosphate GPP , an isoprenoid compound that provides the monoterpene fraction needed for cannabinoid biosynthesis \[ 40 \]. The GPP biosynthesis occurs in the plastid matrix, where it can freely move within the hydrophobic membrane \[ 62 \]. The final step in cannabinoid biosynthesis occurs on the cell wall surface in the extracellular storage lumen \[ 67 \]. Other cannabinoids can be synthesized through isomerization from THC, CBD, and CBC, and conversions between cannabinoids can also occur under specific conditions \[ 34 \]. Biosynthesis, trafficking, and secretion of cannabinoids in Cannabis glandular trichomes GTs. A Structural diagram of secretory GTs. B The distribution of plastids in secretory-stage GT disc cell. Plastids move between cells via apertures in the GT cell wall, aggregating below the surface wall and storage cavity. CBGA likely partitions into the plastid membrane bilayer owing to its lipophilicity. Endoplasmic reticulum ER membrane contact between the ER and plastids may facilitate lipid metabolite transfer through lipid-binding proteins or transient hemifusions. Cannabinoids form lipophilic metabolite droplets within the apical cell wall, creating storage cavity space. With advancements in sequencing technology, multiple versions of the Cannabis genome have been published, providing a deeper understanding of the synthesis mechanism of THC and CBD from CBGA at the genomic level. Grassa et al. However, de Meijer et al. This observation, along with the identification of major quantitative trait loci, suggested the presence of a single locus that exhibits codominant Mendelian inheritance patterns \[ 73 \]. However, Kojoma et al. Whole-genome resequencing of Cannabis revealed that nearly all drug Cannabis samples contained the complete coding sequence of THCAS and two CBDAS pseudogenes, while the majority of fiber Cannabis samples only had the full CBDAS-encoding gene \[ 35 \], suggesting that both genes were initially present and functional in the ancestral state. Early domestication involved polymorphism and random loss of function in one of the genes, with THCAS and CBDAS losing their functions in hemp and in drug Cannabis , respectively, after strong artificial selection. The competitive relationship between these two genes in cannabinoid synthesis likely contributed to their loss of function, which was associated with the increase and decrease in THC content in drug Cannabis and in industrial hemp, respectively. Although the de novo sequencing and resequencing of the Cannabis genomes provided insights into the genetic basis and localization of genes involved in cannabinoid biosynthesis, further research is needed to explore the genomic evolution, biosynthesis, and regulation of target compounds. A deeper understanding at the chromosomal level as well as genome-wide association studies GWAS will contribute to a more comprehensive understanding of these aspects. Manipulation of external environmental factors is a significant area for increasing the cannabinoid content like CBD and decreasing the content of THC. Exogenous hormone treatments, including gibberellic acid, methyl jasmonate, and salicylic acid, influence cannabinoid synthesis in Cannabis \[ 75—79 \]. Additionally, the use of mevinolin can effectively reduce THC content by inhibiting the MEP and mevalonate pathways \[ 75 \]. Furthermore, light conditions differently affect the synthesis of different cannabinoids \[ 79 , 82 , 83 \]. These findings highlight the potential of manipulating external factors to modulate cannabinoid composition in Cannabis. Additionally, transcription factors TFs have been identified as key regulators that activate or inhibit the biosynthesis of plant natural products, effectively enhancing the synthesis of desired secondary metabolites. Numerous reports have highlighted the significance of TFs in regulating secondary metabolite synthesis \[ 84—87 \]. However, the lack of a robust genetic transformation system hinders comprehensive studies on the mechanisms regulating cannabinoid biosynthesis. Plant genetic transformation allows plants to acquire new traits, contributing significantly to variety improvement and gene function elucidation. However, for Cannabis as a regenerative and genetically recalcitrant plant, it is difficult to develop an efficient genetic transformation system, with low regeneration efficiency influenced by factors such as variety, tissue type, plant age, and growth regulator combination \[ 92 , 93 \]. Previous studies employed different methods, such as Agrobacterium -mediated transformation, virus-induced gene silencing, and nanomaterial translocation, to achieve transient expression in Cannabis leaves, albeit with limited efficiency \[ 94—99 \]. Recently, efforts have been made to develop stable genetic transformation systems for Cannabis. Wahby et al. Therefore, it is not feasible to study the cannabinoid biosynthesis regulation through transgenic hairy roots. Zhang et al. Nevertheless, the establishment of an efficient transgenic system, for both transient and stable transformation, remains a significant challenge in Cannabis research, particularly for investigating cannabinoid synthesis regulation and other important traits. GTs arise from the differentiation of epidermal cells and are prominent on the surface of numerous plant species \[ \]. GTs are important defense organs against environmental stress, while also offering significant economic and practical value through their specialized metabolites \[ — \]. Secretory GTs are specifically involved in the synthesis, accumulation, and release of a wide range of metabolites, including organic acids, polysaccharides, polyphenols, flavonoids, alkaloids, and terpenoids \[ — \]. In contrast, nonsecretory GTs primarily function as protective structures without chemical compound secretion \[ \]. Secretory GTs are abundantly present on the tissue surface of multiple plants such as Solanaceae, Labiatae and Asteraceae. Given their similar structure, the secretory GTs may have similar developmental events in different plants species \[ \]. Arabidopsis thaliana only possesses nonsecretory GTs \[ , \]. Consequently, although more comprehensive studies on the morphological aspects, density, and developmental processes of GTs have been conducted on Arabidopsis , the mechanisms underlying the development and specialized metabolite biosynthesis in secretory GTs may not be fully elucidated by using the model plant A. On the contrary, the recently extensive studies in Artemisia annua and Solanum lycopersicum exemplify the transcription factors regulating secretory GTs development. Therefore, we summarized a transcriptional regulatory network of GTs development in these both species in the later part of this review to provide a reference for the regulation of GTs in Cannabis Fig. GTs in Cannabis and their transcriptional regulatory network in Solanum lycopersicum and Artemisia annua. A Scanning electron microscopy analysis illustrating the different GT types on the surface of various tissues in Cannabis sativa. B Summarized transcriptional regulatory network of GT initiation in S. Different shapes represent distinct regulatory factors. Arrow-headed lines indicate upregulation, whereas blunted lines indicate downregulation or inhibition. Cannabis GTs are distinctive secretory structures that are crucial in the production and reservation of cannabinoids, primarily found on the bracts and flowers of female Cannabis plants Fig. In Cannabis , the density of GTs per unit area is generally higher in female plants than that in male ones \[ \]. Female plants exhibit a greater abundance of GTs on bracts, flowers, and other plant parts Fig. Stalked GTs consist of a stalked base with 12—16 secretory disc cells; sessile GTs possess a short-stalked base with eight secretory disc cells; and bulbous GTs are the smallest in size and consist of a short-stalk base with a bulbous head. The type and density of secretory GTs exhibit variations during development. In the early developmental stages in bract and flower, all GT types are present, whereas in the later stages stalked GTs are predominant, indicating that sessile GTs may represent an early stage in stalked GT development \[ 5 \]. The synthesis and storage of cannabinoids primarily occur in secretory GTs \[ \]. Through the isolation and analysis of Cannabis GTs, Happyana et al. Autofluorescence studies further supported these findings and revealed the higher accumulation of monoterpenes in the stalked GTs \[ 5 \]. The analysis of Cannabis GT wall components revealed the presence of loosely bound xyloglucan and pectin polysaccharides. Within the cell wall, the interaction between polysaccharide emulsions and metabolite droplets creates a crucial microenvironment where the boundary between metabolites and polysaccharides acts as a hydrophobic—hydrophilic interface \[ \]. Cannabinoid synthases have greater metabolic efficiency in hydrophobic environments than that in aqueous ones \[ \]. This metabolite—polysaccharide wall boundary may be a favorable microenvironment that is well suited for THCA biosynthesis, which could explain the abundant synthesis and secretion of cannabinoids in GTs \[ \]. Despite their small size, the ability of these GTs to produce and secrete significant quantities of secondary metabolites remains a mystery. Unraveling the mechanisms behind their metabolite synthesis should provide valuable insights for improving the production capacity of natural products in microorganisms and plant chassis. These supercells exhibit extensive cytoplasmic bridges across their cell walls and a polarized distribution of organelles adjacent to the apical surface, which is responsible for metabolite secretion. These supercells are organized as polarized cell syncytia, representing a synchronized fusion of multiple cells with no discernible cytoplasmic connection to the underlying tissue. This polarized cell syncytium configuration presents a potential mechanism for preventing the backflow of metabolites from the trichomes by physically isolating the syncytium from the surrounding tissue. The regulatory mechanisms governing the GT development in Cannabis are not yet fully understood. However, the MIXTA gene, an MYB family transcription factor, has been identified as a regulator of multicellular epidermal hairs and conical cells in the leaves of Antirrhinum majus \[ , \]. To date, the regulatory mechanisms underlying the development of secretory GTs in S. Several TFs involved in the development and morphology of GTs in tomatoes have been identified. Chang et al. Transgenic experiments showed that the hair-absent phenotype was caused by deletion of the entire coding region of Hair. Further, Chun et al. Yu et al. Knockdown of SlMYC1 led to smaller and less dense trichomes, whereas its knockout resulted in the disappearance of secretory GTs. Furthermore, Chen et al. Overall, these tomato TFs related to GT development have provided valuable insights into the genetic mechanisms underlying the development and regulation of GTs in Cannabis Fig. The World Health Organization recommends artemisinin as an essential component of standard antimalarial therapy, with A. Artemisinin is specifically distributed in the secretory GTs in A. Overexpression of miR leads to significant inhibition of GT initiation and artemisinin biosynthesis \[ \]. Conversely, inhibition of miR expression had the opposite effect, indicating that miR negatively regulates the GT initiation and artemisinin biosynthesis. Furthermore, the regulatory mechanisms of candidate TFs related to cannabinoid biosynthesis and secretory GT initiation need to be further elucidated by in vivo and in vitro biochemical strategies. Furthermore, single-cell transcriptome sequencing technology could be used to identify genes specifically expressed in Cannabis secretory GTs, providing an important reference for precise mining of regulatory genes related to Cannabis secretory GT development and cannabinoid biosynthesis \[ , \]. However, the study and medicinal use of cannabinoids have faced challenges owing to legal restrictions on Cannabis and the low natural abundance of most cannabinoids. In addition, the complex structures of cannabinoids hinder their large-scale chemical synthesis. Green bioproduction of cannabinoids based on the theory of synthetic biology could efficiently overcome these limitations by providing a comprehensive understanding of the cannabinoid synthesis pathway, synthesizing and assembling the essential components in microbial cells, and optimizing the fermentation production of target cannabinoids. Therefore, achieving high levels of OA production is crucial for effective metabolic engineering of cannabinoids. Escherichia coli is widely recognized as an ideal host for microbial production owing to its straightforward genetic background and rapid reproduction \[ \]. Tan et al. This finding presents a novel and alternative method for the microbial production of cannabinoid precursors, bypassing the need for the OLS and OAC enzymes derived from Cannabis \[ \]. Yarrowia lipolytica , a safe and lipid-rich yeast, has been successfully engineered to produce various valuable natural products \[ , \]. Introduction of LvaE from Pseudomonas sp. These modifications effectively redirect the carbon flux toward OA production. The optimized protocols led to a remarkable fold increase in OA production, with a titer of 9. In contrast to fungi and plants, amoebae represent a unique group of eukaryotes that express terpene synthases, indicating the presence of an active GPP biosynthesis pathway \[ \]. Reimer et al. Subsequently, the optimization of reaction conditions in D. De novo and high-yield production of cannabinoids in microorganisms or heterogenous plants have been gaining attention for its global demand. Luo et al. Although this yield is only at the milligram level, it demonstrates the potential of Saccharomyces cerevisiae as a microbial chassis for cannabinoid synthesis. Qiu et al. NphB, a prenyltransferase from Streptomyces sp. Therefore, Lim et al. The cannabinoid synthesis titer in yeast was mainly restricted by the enzymatic activities related to the OA pathway; there is thus a need for further exploitation of the modification and optimization of these enzymes. Nicotiana benthamiana is a potential plant chassis to efficiently synthesize cannabinoids. Thies et al. However, OA and its glycosylation products were detected owing to the presence of endogenous glycosyltransferase in engineered N. Reddy et al. Owing to the endogenous modifications of cannabinoid intermediates in tobacco, the chassis may be further directly engineered, especially the gene silencing or knockdown of modified enzymes, to make it applicable for cannabinoid bioproduction \[ \]. Microbial production offers an alternative to natural extraction for prenylated cannabinoids but has challenges such as carbon flux diversion, product toxicity, and GPP essentiality, which can hinder large-scale production. Valliere et al. To enhance enzyme stability, a flow system for CBGA capture was implemented, leading to a titer of 1. The cell-free system was further optimized and designed, optimizing the number of enzymes to only 12 enzymes to produce CBGA, and using the relatively inexpensive substrate acetyl-CoA. These solutions were rapidly achieved through iterative design-build-test cycles, surpassing previously reported results using living cells. However, the large-scale construction of highly complex systems involving numerous enzymes, cofactors, and metabolites using a cell-free system remains a challenge. The utilization of metabolic engineering to produce cannabinoids offers unparalleled advantages compared with both plant extracts and chemical synthesis. Although significant efforts have been made to produce cannabinoids using various chassis cells such as E. Considerable efforts are still required to enhance synthetic efficiency, including the improvement of metabolic flux into the cannabinoid biosynthesis in the chassis, exploration and modification of efficient synthetic enzymes, and optimization of fermentation processes. With the global changes in the legalization of Cannabis , further expansion in the market for green cannabinoid bioproduction is expected. Cannabis , an economically important crop, is restricted or banned for cultivation in many countries owing to it containing the psychoactive compound THC. However, the significant medicinal value of various cannabinoids, such as CBD, found in Cannabis has led to increased demand for cannabinoids, accelerating the legalization of industrial Cannabis cultivation globally. Nevertheless, the lack of comprehensive research on Cannabis GT development and cannabinoid biosynthesis regulation poses major challenges for high CBD and low THC Cannabis varieties breeding and efficient green bioproduction of cannabinoids. To address these issues and provide crucial support for the globalization of the Cannabis industry, there is an urgent need for innovative interdisciplinary and technological research on Cannabis. Recently, metabolic engineering of cannabinoids and the cultivation of new medicinal Cannabis varieties with high CBD and low THC content have gained importance. Cannabinoid biosynthetic genes have been fully elucidated, and they present trichome-specific accumulation. However, the mechanisms regulating cannabinoid biosynthesis and secretory GTs remain largely unclear. Although the regulation of secretory GT development in tomato and A. With the advancements in sequencing technologies, multi-omics data for Cannabis , including the T2T genome, 3D genome architecture and epigenomes for flower or trichomes, single-cell transcriptomes, and transcriptomes and metabolomes of different tissues, will provide important genetic resources for the selection of candidate TFs and noncoding RNAs related to cannabinoid biosynthesis regulation and secretory GT initiation \[ , \]. However, conventional breeding often involves lengthy selection cycles and is less efficient \[ 71 \]. In contrast, molecular breeding offers a more efficient and precise approach, thereby accelerating the breeding process \[ \]. Herbgenomics lays the foundation for molecular genetics of medicinal plants. The integration of multi-omics technology has provided additional genetic information for molecular breeding in Cannabis. Combining multiple published versions of the Cannabis genomes with multi-omics approaches with the aid of artificial intelligence-based data analysis tools has significantly improved the efficiency with which the genetic and molecular mechanisms regulating cannabinoid biosynthesis can be uncovered. Particularly, the relationship between the expression of key enzymes involved in cannabinoid biosynthesis and the THC:CBD ratio and cannabinoid content can be better understood at the genetic and molecular levels. Other breeding strategies such as GWAS use natural populations to identify molecular markers associated with important traits in Cannabis \[ \]. Moreover, GWAS enables the discovery of molecular markers and facilitates the identification of loci and candidate genes related to crucial agronomic traits such as CBD and THC biosynthesis, sexual development, and cellulose quality \[ , \]. Constructing a genetic linkage map of Cannabis using molecular marker technology, leveraging the genetic information from diverse genetic resources, and developing molecular markers for desirable traits will further advance the process of Cannabis breeding \[ , \]. Although significant progress has been made in the de novo synthesis of cannabinoids in yeast, achieving high yields of cannabinoids is still a major challenge. The rate-limiting steps and metabolic bottlenecks in cannabinoid biosynthetic pathways within chassis cells need to be further addressed and optimized to improve the overall efficiency and yield of cannabinoid production. Site-directed mutations of crucial rate-limiting enzymes in cannabinoid biosynthesis could efficiently improve the yields in chassis cells. Cannabinoid accumulation via parallel evolution has been discovered in Helichrysum umbraculigerum , providing a set of alternative enzymes for the synthetic biology of cannabinoids \[ , \]. Further studies can explore the construction of fusion proteins from different plant sources to enhance catalytic efficiency. In addition to microbial and plant chassis, OA synthesis has also been reported in D. Amoebas offer a wider and larger gene repertoire for polyketide and terpenoid biosynthesis, presenting a new approach for the synthetic chassis of cannabinoids \[ \]. In recent times, research on efficient Cannabis cultivation has gained significant attention owing to the substantial market demand. Dioecious Cannabis offers several advantages as a robust model research system for understanding the development of secretory GTs. Cannabis has a relatively short growth cycle of approximately three months, and short-day conditions can induce Cannabis flowering initiation. The generation time can be shortened within nine weeks by controlling light condition \[ \]. Here, we construct a schematic to describe the Cannabis growth cycle and corresponding biotechnologically experimental timeline Fig. Briefly, multiple methods developed and exploited for the genetic transformation of Cannabis generally occur at or before seedlings. Fast cutting propagation usually takes place in the rapid growth stage before inflorescence \[ \]. To avoid genetic heterozygosity introduced by hybrid breeding, as Cannabis is dioecious, female plants can be transformed to produce male flowers by treating with hormones or AgNO 3 \[ \]. The establishment of a stable genetic transformation system in Cannabis deepens our understanding of regulatory mechanisms and developmental biology and is critical in establishing Cannabis as a model plant. Genetic improvement to obtain new Cannabis varieties with superior traits serves as a valuable resource for the medicinal and health utilization of cannabinoids. Cannabis growth and experimental timelines. Cannabis exhibits a relatively short growth cycle, lasting 2—5 months. The seedling stage takes approximately 3—10 days after germination. Subsequently, it enters the vegetative growth stage characterized by slow growth and nutrient absorption focused on root development, lasting for approximately 20—30 days. The subsequently rapid growth of the aerial part is maintained for 10—30 days. Preflowering shoots as cuttings are suitable for asexual propagation. Callus induction and genetic transformation can be performed by taking explants from the hypocotyledonary axis, cotyledon, and tender leaf. The flowering initiation in Cannabis can be induced by subjecting the plants to short-day conditions. After approximately a week, flowers begin to appear at the top of the branches. The flowering phase typically lasts for a month. When treated with hormones or AgNO 3 , female plants can be transformed into male plants. The harvesting of inflorescences for cannabinoid extraction can be performed once the pistils have withered. If female flowers are pollinated during the flowering phase, seeds can be collected after approximately a month. New varieties with high CBD yield can be obtained through hybridization breeding or molecular breeding techniques. In conclusion, Cannabis , an ancient medicinal plant with a longstanding history of global usage over millennia, can make a substantial transformative effect on human health in the future. All authors read and approved the final manuscript. This section collects any data citations, data availability statements, or supplementary materials included in this article. As a library, NLM provides access to scientific literature. Hortic Res. Find articles by Ziyan Xie. Find articles by Yaolei Mi. Find articles by Lingzhe Kong. Find articles by Maolun Gao. Find articles by Shanshan Chen. Find articles by Weiqiang Chen. Find articles by Xiangxiao Meng. Find articles by Wei Sun. Find articles by Shilin Chen. Find articles by Zhichao Xu. These authors contributed equally to this article. Open in a new tab. Click here for additional data file. Similar articles. Add to Collections. Create a new collection. Add to an existing collection. Choose a collection Unable to load your collection due to an error Please try again. Add Cancel. Used a short-chain acyl-CoA synthetase co-expressed with the acetyl-CoA carboxylase. Ma et al. Okorafor et al. Zirpel et al. Introduced and modified more than 15 genes from different species into yeast. Cannabigerolic acid CBGA , olivetolic acid glucoside and cannabigerolic acid glucoside. Gulck et al. Lee et al. Lim et al.

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