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Cannabis sativa L. Recent legal cannabis production in North America and Europe has been accompanied by an increase in reported findings for optimization of naturally occurring and synthetic cannabinoid production. Ultraviolet UV radiation shows potential in stimulating cannabinoid biosynthesis in cannabis trichomes and pre-harvest or post-harvest UV treatment merits further exploration to determine if plant secondary metabolite accumulation could be enhanced in this manner. Well-designed experiments with light wavelengths other than blue and red light will provide more insight into light-dependent regulatory and molecular pathways in cannabis. Lighting strategies such as subcanopy lighting and varied light spectra at different developmental stages can lower energy consumption and optimize cannabis PSM production. Although evidence demonstrates that secondary metabolites in cannabis may be modulated by the light spectrum like other plant species, several questions remain for cannabinoid production pathways in this fast-paced and growing industry. In summarizing recent research progress on light spectra and secondary metabolites in cannabis, along with pertinent light responses in model plant species, future research directions are presented. Secondary metabolites from plants, animals, and microorganisms drive many medical and pharmacological applications, building on thousands of years of traditional medicine Stojanoski, In depth characterization of isolated plant secondary metabolites PSM for medical treatment started at least years ago, and it has progressed exponentially during the last 30 to 40 years Okada et al. One notable and historical medical application is the isolation of morphine from poppy Papaver somniferum seed oil in the early s Krishnamurti and Rao, This alkaloid and its derivatives, opiates, are used for managing pain, yet they have contributed to a deadly and costly opioid crisis because of their addictive nature Dasgupta et al. The cannabis plant Cannabis sativa L. Research on cannabis PSM has grown rapidly because of therapeutic potential. Prior to cannabis legalization, our knowledge of cannabis PSM production primarily stemmed from illegal production operations Vanhove et al. Over the last few years, enormous progress has been made toward advancing cannabis-related medicine Hutchison et al. Contemporary medicine highlights cannabinoids, terpenes, and flavonoids as promising PSMs for treating multiple ailments Aliferis and Bernard-Perron, Evidence suggests that growing conditions i. Light triggers plant secondary metabolism and PSM accumulation, although how optical and spectral properties i. This review aims to bridge the gap between light properties and cannabis PSM production, by recalling PSM origin and function in plants. Available light study findings on cannabis PSM production in response to different light treatments are summarized, with an emphasis on ultraviolet UV radiation during plant growth. PSM are assembled from primary metabolite precursors Seca and Pinto, These PSM are not essential to plants' survival; rather, they allow plants to withstand abiotic and biotic stress drought or water stress, light or predatory stress Bourgaud et al. PSM molecular pathways are conserved between plant families through gene clusters. PSM likely evolved in an environment where biotic stressors played a lesser role in driving evolutive adaptation Tossi et al. Plants evolved mechanisms to protect against this radiation stress by accumulating phenolic and terpenoid compounds that absorb UV radiation and acted as sunscreen in leaves Rozema et al. This allowed photosynthetic organisms to grow in new ecological niches, while exposing themselves to increasing UV radiation Tossi et al. This theory is supported by the apparition of a highly conserved receptor, UV-B Resistance 8 UVR8 in terrestrial plants that mediates plant photomorphogenesis in response to UV radiation Jenkins, ; Tossi et al. Parallel to the abiotic stress response, the large diversity of PSM can also be explained by exposure to biotic stress and the co-evolution of insects and plants during terrestrialization in the Neoproterozoic era 1, to million years ago Theis and Lerdau, ; Labandeira, Plants evolved attractant and deterrent cues through their PSM to favor pollination and decrease predation Kessler and Halitschke, Studies report that cannabis PSM extracts, specifically hemp extracts, effectively repel insects Mcpartland, ; Benelli et al. The cannabis microbiome also influences plant metabolism. A recent review highlights promising avenues of PSM modulation in cannabis through endophytes Taghinasab and Jabaji, Trichomes form a large group of plant structures that are uni- or multicellular epidermal appendages, classified by their origin, form, function, and secretion Werker, These structures are responsible for synthesis and storage of cannabinoids and terpenes in C. They protect plants from light stress Lydon et al. Other mechanisms, including water absorption through dew collection, salt secretion, and alluring function, are reported Werker, Glandular trichomes are found on all anatomical plant parts except the hypocotyl and cotyledon, and non-glandular trichomes are found on stems, leaves, petioles, stipules, bract, and tepals Raman et al. Glandular trichome classification relates to morphological traits and composition of the chemical substance secreted Werker, Three types of glandular trichomes in the cannabis plant are described and size-differentiated: capitate-stalked, capitate-sessile, and bulbous trichomes Dayanandan and Kaufman, ; Hammond and Mahlberg, Capitate-stalked trichomes are found exclusively on flowering regions, whereas capitate-sessile and bulbous trichomes are found everywhere except the hypocotyl and cotyledon Raman et al. Capitate-stalked glandular trichomes have more secretory disc cells than other plants and secrete specialized metabolites in the subcuticular oil storage cavity, instead of through pores formed in the cuticle Tissier, ; Huchelmann et al. Excretory cells secrete a resin in a subcuticular cavity Small and Naraine, This resin contains high concentrations of the economically important cannabinoids, with psychoactive and medicinal properties Dayanandan and Kaufman, ; Small and Naraine, Optimal cannabinoid and terpene biosynthesis in glandular trichomes is of paramount importance to bud quality El-Alfy et al. Cannabinoids, also called meroterpenes or terpenophenols, are PSM synthesized by members of the Cannabaceae family, and several other plant species, including Echinacea purpurea, Echinacea angustifolia, Acmella oleracea, Helichrysum umbraculigerum , and Radula marginata Bauer et al. Three C. This differentiation based on cannabinoid content or cannabis cultivars is inadequate, particularly for the medical industry, since it does not reflect or match the therapeutic and medical properties Russo, Changes in PSM biosynthesis during ontological development of cannabis are well-studied, starting with cannabinoid and monoterpene concentrations in flowers in the first weeks of the flowering phase, and ending with almost four times the quantity in a matter of 7 weeks Aizpurua-Olaizola et al. At least cannabinoids and terpenes have been identified Elsohly and Slade, ; Elsohly and Gul, ; Ahmed et al. PSMs are usually extracted from this, as maximal PSM accumulation is often found in glandular trichomes. Other studies have concentrated on determining the role that flavonoids play in cannabis physiology, and how cannabis-specific flavonoids may be exploited Barrett et al. Figure 1 shows the cannabinoid biosynthesis pathway and precursor formation. Primary biosynthesis steps are impacted by UV radiation and blue light Dolzhenko et al. Cannabinoid biosynthesis starts as isopentenyl diphosphate IPP , formed from glyceraldehyde 3-phosphate G3P , and pyruvate in plastids Mcgarvey and Croteau, GPP also acts as a precursor for monoterpene biosynthesis. Figure 1. A simplified overview of cannabinoid and terpene biosynthesis pathways in cannabis Cannabis sativa L. Enzymes are in dashed line box. Enzymes in shaded blue boxes are upregulated by UV radiation and blue light in Lamiaceae. Olivetolic acid OA sets cannabinoid and monoterpene biosynthesis apart. During these steps, cannabinoids are naturally converted from their acid forms during storage or heating decarboxylation as non-enzymatic catalyzed reactions Veress et al. CBCA is most actively synthesized in young cannabis seedlings and can be found in both drug-type and fiber-type cannabis plants, yet its concentration is relatively low compared to other cannabinoids Kushima et al. Involvement of other enzymes such as cytochrome Ps leads to more complex terpenes diterpenes, C 20 Grof, ; Booth and Bohlmann, Independent of the inflorescence stage, major monoterpenes found in indoor-grown C. As inflorescence matures, monoterpene accumulation increases relative to sesquiterpenes Figure 1 Booth et al. Although more than terpenes have been identified in C. Since robust analytical standards are lacking, reported terpene profiles in C. Elucidation of the underlying mechanisms surrounding terpene biosynthesis in cannabis plants may lead to further exploration and different medical applications for this PSM group Aliferis and Bernard-Perron, Terpenoids a modified class of terpenes with different functional groups are by far the most diverse group, with at least 80, different compounds Christianson, ; Zhou and Pichersky, In recent years, cannabis terpenoids have slowly gained interest Arena et al. However, terpene composition in cannabis resin is dependent upon genetic, environmental, and developmental factors, and highly variable terpene profiles additionally exist between individual plants Fischedick et al. Terpene diversity in cannabis resin is responsible for scent and flavor qualities of cannabis flowers Booth et al. Members of the phenol family, flavonoids, form an important PSM group that aids in the plant's responses to sunlight and UV radiation Downey et al. More than 20 flavonoid types in C. Others, such as cannflavins A, B, and C, are uniquely found in cannabis Barrett et al. Cannabis-specific flavonoids show promising therapeutic effects because of their anti-inflammatory activities Barrett et al. Cannabis-specific flavonoid biosynthesis is not well-established. Figure 2 shows the proposed biosynthetic pathway s for cannflavin A and B in C. Flores-Sanchez and Verpoorte, b ; Rea et al. Although it has not been reported in C. Figure 2. Enzymes in shaded blue boxes are upregulated by UV radiation in Arabidopsis thaliana. Dashed arrows represent proposed enzymatic reactions. O-methyltransferase 21; CsPT3, C. Unlike cannabinoids and terpenes, flavonoid spatial and temporal distribution in cannabis plants does not follow the same pattern Aizpurua-Olaizola et al. Rather, higher flavonoid content is reported in C. Low flavonoid content in cannabis oil and seeds is reported Frassinetti et al. Recent studies show that flavonoid accumulation in inflorescence is variety-dependent and could be an indicator of the susceptibility of the variety to oxidative stress Pavlovic et al. Pavlovic et al. Harnessing the radical scavenger activity and screening ability of flavonoids against UV radiation is a promising means of increasing flavonoid production in medical varieties Agati and Tattini, Plants respond to light stress by producing and accumulating PSM Thirumurugan et al. However, the specific effects of light, including light properties wavelength and intensity and fixture configuration i. These studies primarily focused on PSM accumulation in leaves rather than floral biomass. Table 1 summarizes available studies aimed at determining the impact of light spectrum and lighting configurations on cannabinoid and terpene accumulation. Table 1. A comparison of cannabis PSM yield data compiled with overhead, subcanopy, or supplemental lighting. Different wavelength ranges in UV radiation result in varying cannabinoid accumulation Lydon et al. It has been nearly four decades since the first study suggesting that UV-B — nm radiation affects cannabinoid accumulation in cannabis plants Lydon et al. When the daily dosage of UV-B radiation increased from 0 to It was further noted that UV-B radiation increases trichome numbers. Altitude may be equally important. Notably, UV radiation sources used in both studies had relatively broad spectra, compared to electrical UV radiation sources, such UV-discharge lamps and light-emitting diodes LEDs. Clearly, more studies are required to clarify the impact of UV radiation on cannabis PSM accumulation. The impact of visible light on cannabis PSM accumulation has been investigated with different lighting configurations and different wavelengths Hawley, ; Magagnini et al. A high percentage of blue light cause increased cannabinoid content in cannabis inflorescence drug-type cannabis, high amount of THC Hawley, ; Namdar et al. Subcanopy BGR lighting had a higher impact on terpene accumulation than BR lighting, on both upper and lower canopies Hawley, Conflicting results on the interactive effects between blue light and cannabinoid content, however, were reported recently in fiber-type cannabis hemp Wei et al. Westmoreland et al. Wei et al. As such, it is unknown that whether such variation on the interactive effect between spectral quality and cannabinoid accumulation is caused by light levels or cannabis chemotypes. No physiological theories explain how supplemental green light induces cannabis PSM accumulation. Clearly, both spectral properties and cannabis chemotype used highly impact cannabinoid accumulation, and further investigation on the links between spectral properties, cannabis chemotype, and photoreceptor is required to clarify the spectral effects. Light regimes are elemental to C. Although the studies on cannabis growth and photobiology has expanded in the last few years, a comprehensive review by Aliferis and Bernard-Perron concludes that how light spectra influence cannabis metabolomics is still largely unknown. In particular, how cannabis PSM biosynthesis is impacted by monochromatic light requires further investigation, as most studies to date were conducted under mixed wavelength or full-spectrum light conditions. Figure 3 summarizes what is known of wavelengths and corresponding C. PSM responses. Few studies have attempted to identify the regulatory elements of PSM biosynthetic pathway in cannabis plants Marks et al. Follow up studies are required to place these cannabis proteins in the cannabinoid and flavonoid metabolic pathways. As for terpenes, although several studies indicate that UV-B radiation effects higher monoterpene content in plants that contain glandular trichomes Johnson et al. Figure 3. The impact of wavelengths on Cannabis sativa L. It has been proposed that light-dependent reactions for photosynthesis occur and supply energy for metabolic activity in tomato Solanum lycopersicum type VI glandular trichomes Balcke et al. Using this as a precedence, it may be of interest to evaluate the global carbon and energy balance in C. Although Veress et al. How visible light affects terpene biosynthesis remains elusive due to limited studies and terpene diversity monoterpenes, sesquiterpene, and diterpenes. Drawing from previous studies of other crops may provide some insight and future direction for cannabis terpene production Kessler and Kalske, It was concluded that blue light could generally promote monoterpene content in P. A nascent legal industry with proprietary value slows access to reliable information on indoor cannabis production, postharvest practices and processing of cannabis and cannabis-derived products. Apart from controlling environment light, temperature, nutrients, microbiome etc. For example, UV radiation could be used at the end of the flowering stage or before harvest to increase PSM production. More studies on how light can be manipulated during plant production and post-harvest for consistent PSM production and accumulation are anticipated. Here we review known aspects of photobiology that are relevant to PSM production in C. It is clear that light spectra can be manipulated to target specific cannabis PSM accumulation in different cannabis tissues leaves and buds , resulting in altered potencies. Practically applied, optimized light regimes should reduce necessary electrical inputs while increasing cannabis PSM yields and quality. UV radiation is a powerful tool for stimulating cannabinoid biosynthesis in cannabis trichomes, while visible light alone impacts specific cannabinoid biosynthesis pathways and PSM profiles. UV radiation impacts terpene biosynthesis in other model plants, and this could be useful for cannabis plants. We expect that UV and blue LEDs will be increasingly used to stimulate desirable cannabis PSMs, as they have been widely applied and tailored to other high-value crops. The majority of cannabis studies are conducted under blue- and red-light mixtures, leaving a large sum of wavelengths in the visible spectrum untouched. Gene regulatory and molecular pathways affecting cannabis metabolomics under monochromatic light remain elusive. Lighting strategies such as subcanopy lighting and varying light spectra for different plant growing stages and plant architecture can lower energy consumption and optimize cannabis PSM production, eventually improving the precision of cannabis PSM production, as well as therapeutic capacities. Based on research reviewed, a few experimental directions are proposed to bridge knowledge gaps in cannabis lighting and PSM accumulation research: 1 The impact of narrow-spectrum light on cannabis PSM accumulation. Light spectrum greatly impacts cannabis PSM accumulation, yet there is minimal research available on the impact of narrow-spectrum light as most studies were conducted under either dichromatic or full-spectrum lighting. UV LED sources with different wavelengths are highly available, and the accessibility to both researchers and producers make results more accessible. VD and B-SW led the writing of this paper. B-SW and SM were the major editors. ML is the correspondence point person. All authors contributed to the article and approved the submitted version. The funder was not involved in the study design, collection, analysis, interpretation of data, the writing of this article or the decision to submit for publication. Abrams, D. Should oncologists recommend cannabis? Options Oncol. Agati, G. Multiple functional roles of flavonoids in photoprotection. New Phytol. Ahmed, S. Minor oxygenated cannabinoids from high potency Cannabis sativa L. Phytochemistry , — Aizpurua-Olaizola, O. 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Introduction Secondary metabolites from plants, animals, and microorganisms drive many medical and pharmacological applications, building on thousands of years of traditional medicine Stojanoski, Trichomes And Cannabis Profiling Trichomes Trichomes form a large group of plant structures that are uni- or multicellular epidermal appendages, classified by their origin, form, function, and secretion Werker, Cannabinoids and Cannabis Profiling Cannabinoids, also called meroterpenes or terpenophenols, are PSM synthesized by members of the Cannabaceae family, and several other plant species, including Echinacea purpurea, Echinacea angustifolia, Acmella oleracea, Helichrysum umbraculigerum , and Radula marginata Bauer et al. Cannabis PSMs and Biosynthesis Changes in PSM biosynthesis during ontological development of cannabis are well-studied, starting with cannabinoid and monoterpene concentrations in flowers in the first weeks of the flowering phase, and ending with almost four times the quantity in a matter of 7 weeks Aizpurua-Olaizola et al. Cannabinoids Figure 1 shows the cannabinoid biosynthesis pathway and precursor formation.

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