Illuminating Cannabis sativa L.: The Power of Light in Enhancing C. sativa Growth and Secondary Metabolite Production
1Department of Plant Medicals, Andong National University, Andong 36729, Republic of Korea; smvahsan@gmail.com (S.A.); akhtar.ayoobi@gmail.com (A.A.)
2Department of Applied Biosciences, Kyungpook National University, Daegu 41566, Republic of Korea; injamumrassel@gmail.com (M.I.-U.-H.); shifa.2021@knu.ac.kr (S.S.)
3ABEx Bio-Research Center, East Azampur, Dhaka 1230, Bangladesh
4Department of Agroforestry and Environment, Bangabandhu Sheikh Mujibur Rahman Agricultural University, Gazipur 1706, Bangladesh
5Institute of Genomics for Crop Abiotic Stress Tolerance, Department of Plant and Soil Science, Texas Tech University, Lubbock, TX 79409, USA
6Institute of Cannabis Biotechnology, Andong National University, Andong 36729, Republic of Korea
*Correspondence: atikrahmanbt@gmail.com (M.A.R.); mrahman@bsmrau.edu.bd (M.M.R.); hwchoi@anu.ac.kr (H.W.C.)Abstract
Light is crucial for higher plants, driving photosynthesis and serving as a powerful sensory signal that profoundly modulates growth, development, physiological functions, hormone activation, and biochemical pathways. Various light parameters—quality, intensity, composition, and photoperiod—exert a tremendous influence on plant growth and development, particularly in industrial hemp (Cannabis sativa L.). C. sativa, a crop of historical significance and unparalleled versatility, holds immense value in the food, fiber, and medicinal industries. The cultivation of medicinal cannabis is burgeoning in controlled environments due to evolving healthcare regulations. Optimal light conditions significantly enhance both yield and harvest quality, notably increasing the density of apical inflorescences and the ratio of inflorescence to total aboveground biomass. C. sativa metabolites, especially phenolic and terpene compounds and Phytocannabinoids like CBD (cannabidiol), THC (tetrahydrocannabinol), and CBG (cannabigerol), possess immense medicinal value. Secondary metabolites in C. sativa predominantly accumulate in the trichomes of female flowers and surrounding sugar leaves, underscoring the critical need to boost inflorescence weight and metabolite concentrations while ensuring product consistency. Different light parameters distinctly impact C. sativa’s metabolic profile, providing a robust foundation for understanding the optimal conditions for synthesizing specific secondary metabolites. While the effects of light measurement on various crops are well-established, scientific evidence specifically relating to light quality effects on C. sativa morphology and secondary metabolite accumulation remains scarce. In this review, we critically summarized how different light properties can alter cannabis growth (vegetative and reproductive), physiology and metabolism. Furthermore, the mechanisms by which specific wavelengths influence growth, development, and secondary metabolite biosynthesis in C. sativa are not fully elucidated, which could be a prospective task for future researchers. Our review paves the way for a profound understanding of light’s influence on C. sativa growth and advancements in greenhouse settings to maximize metabolite production for commercial use.
1. Introduction
Cannabis sativa L., a short-day photoperiodic species, is cultivated for an array of diverse purposes encompassing fiber production, food sources, medicinal applications, and recreational consumption [1]. This botanical species is taxonomically bifurcated into C. sativa and drug-type varieties, a classification predicated on their utilization purpose and the distinctive chemical profiles of their inflorescences [1]. In the realm of pharmaceutical applications, C. sativa is predominantly cultivated for its prolific production of secondary metabolites, which include cannabinoids (CBD, CBG, THC, CBDA, THCA, CBGA, etc.) terpenes, and flavonoids [2]. Notably, these bioactive compounds predominantly accumulate within the glandular trichomes located on the female flowers and the proximate sugar leaves. The biosynthetic pathways responsible for the synthesis of these secondary metabolites can be delineated into three principal metabolic routes: the polyketide synthase (PKS) pathway, the mevalonic acid (MVA)–cytosolic mevalonate (MEV) pathway, and the Palstidial methylerythritol phosphate (MEP) pathway [1,3]. Specifically, the PKS pathway is instrumental in the production of Olivetolic acid (OLA), which serves as the precursor molecule for phytocannabinoid biosynthesis (Figure 1) [1]. Within the glandular trichomes, which are predominantly found on the unfertilized female inflorescences, the synthesis and accumulation of cannabinoids and terpenes occur [1,2]. These inflorescences, therefore, represent the most pharmaceutically valuable segment of the plant, owing to their high concentrations of bioactive secondary metabolites. This intricate biosynthetic process underscores the significance of unfertilized female inflorescences in the pharmacological utility of C. sativa [1,2].
The life cycle of industrial C. sativa can be delineated into four distinct stages: (i) germination/cloning, (ii) vegetative growth, (iii) flowering and seed formation, and (iv) senescence [4]. The maximization of marketable biomass production, particularly the mature, unfertilized female inflorescences, constitutes a primary objective in the cultivation of cannabis [1,2]. To propel this nascent industry forward, it is imperative to concentrate research efforts on cultivars exhibiting minimal variability in growth metrics, biomass yield, seed quality, and secondary metabolite profiles [5]. Comprehending the intricate interplay between genotype and environmental variables, including photoperiod, in the phenology of C. sativa varieties is essential for optimizing biomass production, enhancing product potential, and minimizing nutrient inputs to achieve sustainable agricultural practices [5]. Although the concentrations of secondary metabolites are predominantly dictated by plant genetics, environmental conditions exert a significant modulatory influence [5]. Thus, agricultural productivity can be substantially augmented through the meticulous manipulation and optimization of environmental parameters to favor desired yield outputs [5]. Indoor cultivation, in particular, presents a distinctive advantage, facilitating the rigorous control of environmental factors such as light quality and temperature [6,7]. This controlled environment paradigm stands in stark contrast to traditional outdoor farming, where agricultural productivity is frequently hindered by unpredictable climatic conditions [6,7]. Therefore, the strategic optimization of these environmental variables in indoor cultivation settings holds the potential to substantially elevate yield outcomes [6,7].
Light conditions, encompassing light quality, light intensity, and photoperiod, exert a profound influence on plant growth and development, with light quality representing the most intricate factor [8]. Indeed, plant responses to these varied light conditions are mediated by a diverse array of photoreceptors. The light spectrum can, both directly and indirectly, impact photosynthesis by modulating leaf structure, size, senescence rate, photosynthate transport, stomatal conductance, transpiration, gas exchange, chlorophyll content, plant height, stomatal opening, circadian rhythm, photomorphogenesis, flowering time, and chlorophyll biosynthesis [9]. Extensive research has elucidated the impact of light cycles and environmental conditions on plant development from germination to senescence, with numerous studies examining the correlation between photoperiod and flowering patterns across different plant families by regulating flowering-induced genes [10,11]. Light stands as a pivotal environmental factor for plant growth and development, with stressful conditions capable of inducing the excessive production of reactive oxygen species (ROS) (Figure 1) [12,13]. ROS can wreak havoc on cellular components, including carbohydrates, lipids, proteins, and DNA, potentially leading to plant death. To combat this onslaught, plants deploy a sophisticated arsenal of enzymatic and non-enzymatic antioxidants to regulate ROS production [14].
Photosynthetic carbon fixation, which hinges on light energy, is acutely responsive to light intensity at specific wavelengths. Photoperiodism governs plants’ developmental responses to the daily light-dark cycle, while photomorphogenesis orchestrates the profound effects of light quality on plant development and physiology [5,12,13,14]. Secondary metabolism in plants is heavily influenced by external factors such as light intensity, light spectrum, day length, mineral nutrition, plant architecture, and temperature. In C. sativa, secondary metabolites serve critical photoprotective roles. Evidence reveals that light stress or manipulation can dramatically alter the plant’s metabolomic composition. Adjusting wavelength composition can also significantly impact phytohormone activity, stimulating flowering, inhibiting stem elongation, or reducing plant height. Chloroplasts, the epicenters of light sensing, dynamically alter their ultrastructure in response to varying photoperiods. The quality, quantity, and length of light are paramount, influencing the excitation of Photosystem I (PSI) and Photosystem II (PSII), the linchpins of photosynthesis (Figure 1) [5,12,13,14].
Moreover, light parameters can bolster plant resistance to drought and salt stress, and recent evidence underscores the impact of light duration on plant responses to pathogen infection. Light manipulation holds immense potential, capable of regulating various aspects of insect life history, presenting a powerful tool for insect pest management in controlled environments [5,12,13,14]. The yield quantity and quality of inflorescences in C. sativa can be markedly enhanced within controlled environment cultivation systems, where all environmental parameters and cultivation practices are meticulously regulated. Indoor cultivation often becomes indispensable due to unsuitable climatic conditions, stringent regulatory restrictions, or a synergistic effect of both factors [5,15]. Among the myriads of factors influencing successful indoor cannabis cultivation, light—encompassing photoperiod, quality, and intensity—remains paramount [5,15]. To ensure uniform environmental conditions and produce a consistent product, there is a burgeoning trend towards cultivating cannabis in controlled indoor environments. Within these ‘in-door’ facilities, artificial lighting systems provide the requisite light, with fixtures tailored to vary in intensity and spectral composition [5,15].
Cannabis, cultivated for both recreational and medicinal purposes, is recognized as a high-value crop predominantly grown within controlled environment production facilities, such as greenhouses utilizing natural light or growth chambers and vertical farms devoid of natural light [5,15]. These cultivation setups facilitate consistent, year-round production. The propagation, vegetative growth, and flowering stages of indoor cannabis production exhibit distinct photoperiod and light-intensity requirements [5,15]. Optimal post-vegetative stage morphology is contingent upon the production system employed by cultivators, which encompasses variables such as the length of the vegetative stage, planting density, substrate and root zone volume, and the type of trellising system implemented during flowering [5,15]. Nevertheless, the overarching objective is to ensure high transplant success rates and robust vegetative growth. Maintaining rigorously controlled conditions is indispensable for the consistent production of medical cannabis, particularly concerning inflorescence yield and the concentrations of secondary metabolites [5,15]. These controlled environment cultivation systems not only facilitate the optimization of growth conditions but also ensure the production of a high-quality, uniform product, thus meeting the stringent demands of both medicinal and recreational cannabis markets [5,15].
In this review, we conducted a comprehensive examination of recent research focused on the impact of indoor light on the growth, development, and secondary metabolite production in C. sativa, situating our analysis within the broader context of light signaling in plant physiology. Our findings underscore pivotal areas where the intricate interaction between light, and C. sativa necessitates further elucidation to achieve a comprehensive understanding of its underlying mechanisms.
2. Role of Light in Plant Growth, Development, and Secondary Metabolism: A Background Story of Mechanism
Light, a critical environmental factor, influences plant growth, development, and metabolism, functioning in both photosynthetic and signaling capacities for plant morphogenesis [16]. Variations in light quality (spectrum), intensity, and duration catalyze critical physiological and biochemical reactions, dramatically shaping plant form and function (Figure 1) [16]. Plants have evolved a sophisticated circadian clock to synchronize various processes with the light/dark cycle, and metabolic reprogramming in response to light is crucial for optimal plant growth and development [16]. Understanding these processes is pivotal for improving crop production efficiency and agricultural productivity [16].
Light intensity, spectrum (280–800 nm), and day length significantly affect reactive oxygen species (ROS) formation during photosynthetic electron transport (Figure 1), influencing the cellular redox state crucial for metabolic adjustments that enable plant survival under various environmental conditions [17,18]. The dynamic shifts in light conditions, both spatial (latitude, altitude) and temporal (daily, seasonal), are compounded by fluctuations in temperature, water supply, and stress factors. Consequently, light-governed redox control profoundly influences core metabolic pathways (carbon, nitrogen, amino acids, sulfur, lipids, and nucleic acids) and secondary metabolism (terpenoids, flavonoids, alkaloids) (Figure 1) [17,18].
ROS accumulation is particularly intense near electron transport chains, necessitating precise, compartment-specific redox adjustments [17,18,19]. Within chloroplasts, excess ROS emerges near thylakoid membranes during photosynthesis, specifically in the electron transport between photosystems II and I (Figure 1). In mitochondria, ROS accumulate around inner membranes during respiration [17,18,19]. The antioxidant system is crucial in maintaining optimal ROS levels, safeguarding cells from oxidative devastation under various environmental stresses [15,17,18]. Dominant antioxidants include the ascorbate–glutathione cycle, α-tocopherol, carotenoids, flavonoids, thioredoxins (TRXs), peroxiredoxins (PRXs), superoxide dismutase (SOD), catalase (CAT), peroxidase (POD), and glutathione S-transferases (GSTs) [15,17,18]. The synergistic roles of ROS and antioxidants in light-dependent redox regulation are critical for plant adaptation to environmental changes. The strategic metabolic engineering of light signaling intermediates to modulate secondary metabolite accumulation is essential for maximizing plant growth and development [15,17,18] (Figure 1). The light-induced synthesis of photoprotective secondary metabolites is indispensable [17,18,19].
The spectral regulation of metabolism is of utmost importance due to the precise absorption maxima of chlorophylls and the acute sensitivity of photoreceptors (Figure 1). Plants employ sophisticated photoreceptors to precisely adapt, to environmental light cues, drive extensive transcriptome reprogramming, and induce developmental and physiological changes (Figure 1) [16,20]. Light perception, which is crucial for photomorphogenesis and metabolism, involves specific photoreceptors (Figure 1) distinct from photosynthetic pigments that operate independently of photosynthesis [16,20]. Five formidable classes of photoreceptor proteins have been identified in plants: phytochromes, cryptochromes, phototropins, the ZEITLUPE (ZTL)/FLAVIN-BINDING KELCH REPEAT F-BOX 1 (FKF1)/light oxygen voltage (LOV) KELCH PROTEIN 2 (LKP2) complex, and UV RESISTANCE LOCUS8 (UVR8) [16,20]. Phytochromes, which are essential for red and far-red light responses, exist in two states: Pr, absorbing red light (650–670 nm), and Pfr, absorbing far-red light (705–740 nm) [16,20]. Pr converts to the active Pfr upon absorbing red light, and Pfr reverts to Pr upon absorbing far-red light. Phototropins, membrane-bound and activated by blue light, are critical in photosynthesis, regulating chloroplast movement, stomatal dynamics, and optimizing carbon dioxide and water exchange [16,20,21]. The ZTL/FKF1/LKP2 complex, sensitive to blue and UV-A wavelengths, governs circadian rhythms and flowering [16,20,21]. UVR8 receptors, detecting UV-B radiation, mitigate its harmful effects by inducing gene expression linked to antioxidant production. They also regulate vital plant responses, such as stomatal behavior and chlorophyll balance [16,20,21]. These advanced systems allow plants to adjust their growth and development, ensuring survival and optimal performance in dynamic light environments [16,20,21,22,23,24].
Photoreceptor-mediated light perception initiates a signaling cascade that markedly changes gene expression and influences key plant physiological responses [16,20,21]. In Arabidopsis, a staggering 20% of genes are regulated by light, with the bZIP transcription factor ELONGATED HYPOCOTYL 5 (HY5) emerging as a pivotal regulator in light-mediated development and metabolism. HY5 integrates light signals with hormonal and nutrient pathways, while Phytochrome Interacting Factors (PIFs) serve as strong negative regulators of this pathway. The E3 ubiquitin ligase CONSTITUTIVELY PHOTOMORPHOGENIC1 (COP1) controls these transcription factors by targeting them for degradation. HY5’s versatility spans light, hormone, and stress signaling, operating downstream of phytochromes, cryptochromes, and UVR8 to regulate gene expression [16,20,21]. Its interactions with SUPPRESSOR OF PHYA (SPA) proteins and other transcription factor families highlight its central role in light signaling. The transcription of light-regulated genes significantly impacts plant growth, development, and seasonal adaptation [16,20,21]. Hormones such as auxin, ethylene, jasmonic acid, gibberellic acid, and abscisic acid interact with light signaling pathways, influencing both vegetative and reproductive stages [16,20,21,25,26]. Recent advancements have clarified hormone signaling (Figure 1) mechanisms, revealing how light perception profoundly affects hormonal regulation in processes like germination, flowering, and morphogenesis [16,20,21].
The circadian clock aligns plant physiology and development with daily and seasonal environmental changes, controlling processes such as photosynthesis and stomatal movements [17,26]. Chromatin regulation creates rhythmic gene expression networks that are active at specific times and managed by the clock through feedback loops, involving MYB transcription factors (Figure 1) [17,26]. Shade avoidance, which is crucial for survival, is linked to hormone signaling, such as the auxin-mediated elongation of hypocotyls and petioles [17,25,26,27]. Plants detect shading from neighboring vegetation through dramatic shifts in the red-to-far-red (R–FR) light ratio, sensed by phytochromes.
The response cascade involves hormones like gibberellins (GA), ethylene (ET), auxin, Brassinosteroids, cytokinins (CK), and jasmonic acid (JA) (Figure 1) [17,25,26,27]. Auxin plays a pivotal role in elongation phenotypes and its pathway is tightly regulated when sensing nearby plants. Auxin-related genes dominate a significant portion of the shade avoidance transcriptome [17,25,26,27]. The regulation of apical dominance is another critical aspect governed by endogenous auxins and cytokinins. These growth regulators are instrumental in nutrient diversion, gene expression modulation of axillary bud growth, and fine-tuning the auxin/cytokinin ratio (Figure 1) [17,25,26,27]. The synthesis of auxins at the plant apex and their strategic transport to axillary buds play a crucial role in maintaining apical dominance [17,25,26,27]. These mechanisms are key to understanding plant growth and development in response to light signals and underscore plant adaptability and resilience [17,25,26,27].
Synchronized flowering is crucial for plants, particularly in regions with significant seasonal variation and those requiring cross-pollination [26,28]. Because flower and seed development are energy-intensive processes, they must occur under optimal conditions. Therefore, flowering is meticulously regulated by environmental factors such as day length and temperature, as well as internal signals, including hormonal levels, sugars, and plant age (Figure 1) [26]. Phytochromes A and B (Phy A and Phy B) and cryptochrome 2 (cry2) are crucial photoreceptors that mediate photoperiodic flowering [26]. Different hormones further influence this process. Understanding how these environmental cues and hormonal signals interact is key to deciphering the mechanisms that govern the timing of flowering in plants [26].
Light conveys critical environmental information to plants influencing defense hormone signaling, resource allocation, and adaptation [27]. Jasmonate (JA) and salicylic acid (SA) are powerful defensive phytohormones, and their pathways are profoundly influenced by light, impacting plant defense, growth, and development (Figure 1). Photoreceptors and transcription factors in phototransduction are integral to these hormones’ signals [27], significantly advancing the use of artificial lighting for crop growth and disease management in greenhouses. Jasmonate orchestrates the accumulation of vital defense-related metabolites and proteins, along with MYB, and trichome-specific transcription factors that play crucial roles in UV light signaling and terpenoid production (Figure 1). The light spectrum significantly controls terpenoid biosynthesis and determines plant quality, including aroma, flavor, color, and medicinal properties [27]. Light conditions such as blue, red, and far-red light enable HY5 to drive secondary metabolite production. In Artemisia annua, HY5 binds to specific transcription factor genes, dramatically upregulating their expression and fueling polyphenol, terpenoid, and alkaloid biosynthesis [27,28,29,30,31].
Artificial lighting, such as high-pressure sodium (HPS) lamps, light-emitting diodes (LEDs), and fluorescent lamps, are crucial for indoor cannabis cultivation to meet the light energy needs of the plants. Over the past decade, the use of LEDs in horticulture has skyrocketed due to their overwhelming advantages over traditional light sources (Figure 1). LEDs boast unparalleled energy efficiency, minimal heat emission, and exceptional longevity. They offer superior efficacy and significantly lower operating costs compared to HPS lamps [5,16]. They can emit specific wavelengths, enabling groundbreaking research on the effects of different wavelengths on secondary metabolite production, although data on the effects of varying radiation intensities and qualities on cannabis secondary metabolite composition are limited and sometimes contradictory [5,16]. LED adoption has revolutionized horticultural practices by enabling growers to customize the light spectrum and intensity for different crops and developmental stages, thereby enhancing production schedules, crop yield, and quality. Traditionally, high-intensity discharge (HID) lamps like metal halide (MH) and high-pressure sodium (HPS) lamps have dominated greenhouses and growth chambers (Figure 1) [5,16]. However, the exploration of monochromatic LED systems as replacements for traditional light sources is underway, particularly in space greenhouses, to optimize crop production and quality through meticulously designed light recipes. By combining LEDs of various colors, growers can create a tailored light spectrum at the desired intensity, effectively modulating different plant functions [5,16]. This provides a powerful tool for controlling plant growth and photomorphogenesis [5,16]. These advancements in LED technology hold immense potential for enhancing horticultural practices both on Earth and in space. They enable precise control over the light environment to meet the specific needs of plants at various stages of development [5,16]. Matching LED wavelengths to plant photoreceptors can maximize output and modify plant shape and metabolism, making LEDs indispensable for sustainable production and photomorphogenesis research [5,16].
3. Light Effect on Growth and Development of C. sativa
4. Conclusions and Prospects
Light conditions, including intensity, spectrum, and duration, wield an extraordinary influence over morphogenetic responses, playing a pivotal role in regulating primary and secondary metabolism in plants such as C. sativa. Mastering metabolic adjustment is paramount for optimizing plant growth and development while minimizing the devastation of adverse environmental conditions. The intricate redox-mediated effects of light intensity, photoperiod, and spectrum on primary and secondary metabolism within individual cell compartments demand rigorous investigation, charting critical directions for future research. Light serves as a highly promising abiotic elicitor for stimulating the production of vital metabolites in various in vitro plant systems. Among the diverse abiotic elicitors, light has garnered significant attention due to its precise wavelengths, cost-effectiveness, and long-lasting nature, making it an ideal tool for enhancing secondary metabolite synthesis. Numerous studies have demonstrated that different light sources, with varying qualities and intensities, substantially boost secondary metabolite accumulation across multiple plant species under in vitro conditions. These findings open the door to a deeper exploration of light as a potent elicitor. However, the focus of most studies has been primarily on plant growth and development, as well as primary metabolite generation, underscoring the scarcity of comprehensive insights into the regulatory factors governing light-induced elicitation mechanisms. As a result, many investigations fail to fully elucidate the mechanisms by which light enhances the production of pharmacologically valuable secondary metabolites, as these mechanisms likely vary depending on plant species, culture conditions, and the specific light source applied.
As our comprehension of hormone pathways and photomorphogenic processes deepens, it becomes imperative to unravel the mechanisms through which light signaling pathways intersect with hormone signaling in C. sativa. Groundbreaking advances in light-regulated hormone signaling technologies now empower researchers to delve into the spatiotemporal patterns of signaling as perceived by receptors. The burgeoning understanding of transcriptional and protein regulation presents formidable challenges but is essential for pioneering future progress. Artificial lighting in horticulture has long been a cornerstone for assimilation and photoperiodic functions. However, recent monumental advancements in plant photomorphogenesis and metabolism have heralded the era of state-of-the-art lighting systems and innovative strategies, such as photo-selective greenhouse covers, to manipulate light for supreme control over plant development and metabolism. This revolutionary approach promises to redefine and elevate the optimization of plant growth and productivity through masterful light manipulation. More species and variety-specific trials and research can elucidate the undiscovered molecular mechanism underlying the physiology and metabolism of C. sativa.
Institutional Review Board Statement
Not applicable.
Data Availability Statement
Data will be available upon request.
Conflicts of Interest
The authors declare that they have no competing interests.
| Sl. No. | Variety | Effect of Light (Spectrum, Intensity, Photoperiod) | Duration | Growth Condition | Growth and Developmental Changes | Metabolic Changes | References |
|---|---|---|---|---|---|---|---|
| 1. | WR, CCBD | Red light (600–700 nm), blue light (400–500 nm), far-red light (700–800 nm) | 21 days | Temperature: 25–28 °C RH: 65–85%. | Red and blue light did not significantly affect rooting. Far-red light improved adventitious rooting and stem elongation. | Blue and red lights were not highlighted in this research. Far-red light may induce auxin biosynthesis and carbohydrate content in stem cuttings. | [46] |
| 2. | Yunma 1 | Different light intensities using LED (30 μmol m−2 s−1, 80 μmol m−2 s−1, 130 μmol m−2 s−1, 180 μmol m−2 s−1) | 30 days | Temperature: 24/22 °C day/night RH: 60% Photoperiod: 16 h light/8 h dark | Stem diameter and root dry and fresh weight increased at 130 μmol m−2 s−1. Decreased plant height with increasing light intensity (180 μmol m−2 s−1). | Amino acids, soluble proteins content decreased with increasing LI. Sugar content increased with increasing LI. | [47] |
| 3. | Gelato | Five light-intensity target levels (200, 450, 700, 950, and 1200 μmol m−2 s−1) | 21 days | Daytime temperature and RH: 26 ± 3 °C and 30 ± 9%, 26 ± 3 °C and 30 ± 8%, and 25 ± 2 °C and 30 ± 8%. Nighttime temperature and RH: 23 ± 2 °C and 37 ± 5%, 23 ± 2 °C and 36 ± 5%, and 23 ± 2 °C and 36 ± 5% | Plant height, growth index, nodes, stem thickness, and aboveground dry weight grew quadratically and asymptotically with light LI. PPFD levels between 600 and 900 μmol m−2 s−1 appeared to achieve an appropriate balance in optimizing key morphological parameters. | This experiment found no influence on blue or red light. Photosynthetic capability increased with light intensity. | [48] |
| 4. | Yunma 1 | Different light intensities (30 μmol m−2 s−1, 80 μmol m−2 s−1, 130 μmol m−2 s−1, and 180 μmol m−2 s−1 using LED (red, blue and white) | 30 days | Temperature: 24 °C/22 °C (day/night) RH: 60% Photoperiod: 16 h/8 h | Significant impact on plant development, photosynthesis, and antioxidant enzyme activity in different light intensities. | High light intensity increased C. sativa metabolic profile, lipids, phenolic acids, flavonoids, amino acids, organic acids, alkaloids, nucleotides, sugars, alcohols, vitamins, cannabinoids, coumarins, lignans, terpenoids, tannins, and carbohydrates. | [49] |
| 5. | FINOLA | Red 600–700 nm, far-red 700–800 nm, short wavelength (blue 400–500 nm, UV-A 315–400 nm, and UV-B280–315 nm) radiation | 78 days | Temperature: 24/22 °C day/night RH: 60%/50% day/night | Low R/FR increased plant height and reduced inflorescence production. Short-wavelength radiation did not affect plant morphology or inflorescence yield. | Compared to low R/FR ratio treatment, high R/FR ratio treatment enhanced CBD, THCVA, CBGA, and terpene concentrations. Monoterpene and myrcene concentrations rose with UV-B. No individual light effects were discussed. | [50] |
| 6. | Cultures of C. sativa, accession RTG-XX | LEDs emitted blue (456 nm), red (657 nm), and some white light | -- | Temperature: 27 °C Photoperiod: 18 h, PPFD: 50 μmol m−2 s−1 | Long-term in vitro photosynthesis and respiration in response to various growth conditions. | -- | [51] |
| 7. | Cannabis sativa | Photoperiod | 50 days | Long-day (≥18 h of light) and short-day lighting (≤12 h of light) | Morphological characteristics were enhanced. | [52] | |
| 8. | Bamahuoma | White light (control) (WL), blue light (BL) (450 nm), red light (RL) (650 nm), and 50% blue light with 50% red light (RBL) | 14 days | Temperature: 25/20 °C (day/night), RH: 70–90% | BL boosted the shoot’s fresh and dry biomass, number of leaves/plants, stem diameter, root length, and chlorophyll concentration compared with WL. | BL increases net photosynthesis, stomatal conductance, and transpiration, and decreases lipid peroxidation, superoxide dismutase, and peroxidase activity. RL and RBL significantly reduced the plant biomass and gas exchange parameters with enhanced antioxidant enzyme activities. | [40] |
| 9. | Meridian | PPFD of either 600, 800, or 1000 μmol m−2 s−1 for 12 h day−1 or PPFD of 600 μmol m−2 s−1 plus ultraviolet (UV, 280–400 nm) for either 12 h day−1 of 50 μmol m−2 s−1 from LEDs with a peak wavelength of 385 nm for 45 days (UVA) or 5 h day−1 of 3 μmol m−2 s−1 of wideband ultraviolet fluorescent lighting (UVA + UVB). | 45 days | Temperature and RH: 26 ± 1.2 °C and 40 ± 6.9% (day) Temperature and RH: 22 ± 1.9 °C and 47 ± 3.9% (night) | High LI can substantially increase cannabis yield compared to UV light. Above-ground biomass metrics were 1.3–1.5 times higher in the highest vs. lowest PPFD treatments. | Total foliar THC concentration was unaffected by UV exposure. Sugar leaves (i.e., small leaves associated with inflorescences) of plants in the UVA + UVB treatment had ≈30% higher THC concentrations. High PPFD levels can substantially increase cannabis yield | [53] |
| 10. | Stock plants of ‘Suver Haze’ | Day-length extension lighting with four light intensity treatments of 1.0, 2.5, 5.0, and 10.0 μmol m−2 s−1 PPFD | 24 days | Temperature: 26 °C RH: 85% to 95% Photoperiod: 16 h PPFD: 120 μmol m−2 s−1 | The DE impacts stem length, nodes, and roots. The photoperiod and light intensity affect blooming and inflorescence. | --- | [54] |
| 11. | ‘Gelato-27’, ‘Grace’, and ‘Meridian’ | Blue, red, UV-A, white, and fluorescent LEDs. The photon flux ratios of blue (B; 400–500 nm) and red (R; 600–700 nm) narrowband LED treatment combinations were (1) BR, fixed spectrum of B15/R85; (2) B, B75/R25 on day 0–2 followed by B15/R85 on day 2–14; (3) B+UVA, B75/R25 on day 0–2 followed by B15/R85 on day 2–14 plus 15 μmol m−2 s−1 of UVA on day 7–14; (4) B50, B15/R85 on day 0–7 followed by B50/R50 on day 7–14 | 21 days | Temperature and RH: 26 ± 3 °C; 30% ± 9%, 26 ± 3 °C; 30% ± 8%; 25 ± 2 °C, and 30% ± 8% (day) Temperature and RH: 23 ± 2 °C; 37% ± 5%, 23 ± 2 °C; 36% ± 5%; 23 ± 2 °C, and 36% ± 5% | Blue light increased stem thickness and root index compared to other treatments. No spectrum treatment effects on the percentage of cuttings that rooted, and root index values were higher in cuttings grown under B+UVA vs. F. Relative root dry weights of plugs from the B, B+UVA, B50, and F treatments were higher than the W treatments. Spectrum treatments did not affect the chlorophyll content index. Cuttings under B had thicker stems than BR and W, and those under F showed the lowest percentage of new above-ground growth. | --- | [55] |
| 12. | Finola and USO31 | Plants were grown under white (W) and purple (P) light at different photoperiods (16/8, 20/4, and 24/0) | 35 days | LED light (~80 μmol m−2 s−1, 22 ± 2 °C) | White light yielded the highest PItotal. Purple light yielded the lowest PItotal. | A 16/8 photoperiod, regardless of the light type, yielded the lowest TBARS contents. | [56] |
| 13. | Kanada and E19 | Red and far-red ratio (R/FR) from ceramic metal–halide lamps (CMHs) and high-pressure sodium lamps (HPSs), LEDs | 84 days | Night temperature: 18 °C Day temperature: 23.5°C Humidity: 80% | Modulate the photosynthetic rate | Light spectra significantly influenced CBDA and terpene concentrations | [57] |
| 14. | ‘Babbas Erkle Cookies’ accession | Blue light (430 nm), red light (630 nm), rose light (430 + 630 nm, ratio 1:10), purple light (430 + 630 nm, ratio 2:1), and amber light (595 nm) | 70 days | Temperature: ~28 °C RH: 40–55% (day) Temperature: 25–27 °C, RH: 50–65% (night) Photoperiod: 18 h light (vegetative), 12 h light (flowering) | LED light treatments had lower fresh mean inflorescence mass than the control (HPS, 133.59 g plant−1), and monochromatic blue light yielded the least fresh inflorescence mass (76.39 g plant−1) | Blue light increased the concentration of cannabinoids, including THC and CBG, as well as terpenes. Blue light had a lesser impact on cannabidiol (CBD) biosynthesis. | [58] |
| 15. | Cannatonic, Hindu Kush, and Northern Lights | 10/14 h (light/dark), 12/12 h (light/dark), 14/10 h (light/dark) | 75 days | Temperature: 25 °C RH: 50% | Flower biomass yields were highest for all lines when treatments started with 14L/10D. | A 14L/10D photoperiod significantly decreased THC concentration. Conversely, in Cannatonic, all 14L/10D treatments significantly increased CBD concentration. | [59] |
| 16. | Black Triangle, Garlic Jelly, Ghost Train Haze, Powdered Donuts, Chem de la Chem, Legendary Larry, Gorilla Glue, OG Kush, Incredible Milk, Blue Dream | All treatments were based on a standard 24 h day and included 12 h, 12.5 h, 13 h, 13.5 h, 14 h, and 15 h of light. Approximate light intensity of 360 μmol m−2 s−1 from white LEDs | 3–4 weeks | Temperature: 25.00 ± 0.84 °C RH: 78.40 ± 8.80% | Flowering initiation occurred in all cultivars under all photoperiod treatments up to 14 h. Delays in flowering initiation between 14 h and 12 h varied from 0 to about 4 days, depending on the cultivar. Some cultivars began flowering under 15 h conditions, but floral tissues did not progress beyond the initiation phase. | --- | [60] |
| 17. | --- | Red light (600–700 nm), Green light (500–600 nm), Blue light (400–500 nm) | 14 weeks | Day temperature: 23.2 °C ± 0.1 °C RH: 40.7% ± 0.1% Night temperature: 22.2 °C ± 0.1 °C RH: 41.1% ± 0.3% | Blue light resulted in a higher bud yield than white light. | THC/CBD was unaffected by light. In individual light effects, no phytochemical alterations were identified. | [44] |
| 18. | Finola | Blue and white light | 30 days | Temperature: 20 °C Photoperiod: 16/8 h (light/dark) | White light increased plant shoot and root length. | Leaf pigments were higher under blue light. DPPH, FRAP, flavonoids, and total flavanol content; phenolic acids were not influenced by light type. | [61] |
| 19. | High THCA variety, balanced CBDA/T CA variety, high CBDA variety | White LED, two ratios of blue + red LED (1:4 and 1:1), and a spectrum generated by HPS | 58 days | Temperature: 25 °C, RH: 50% | The highest inflorescence yields occurred when the spectrum was limited to a 1:1 red-to-blue ratio, and in two of the three varieties tested, a 1:4 blue-to-red ratio produced similar results. The lowest yields were observed under white light with a 1:1 blue-to-red ratio. | Blue-rich light-enhanced CBGA accumulation more than far-red-rich HPS light. | [15] |
| 20. | Xinma | CK, high-pressure sodium as light source, (R/B (ratio of red light to blue light) 9.30:1, PPFD (photosynthetic photon flux density) 19:1); LED1 (R/B 9.20:1; PPFD 129); LED2 (R/B 1.61:1; PPFD 540); LED3 (R/B 6.47:1; PPFD 28.2); LED4 (R/B 7.15:1; PPFD 41.7); LED5 (R/B 16.8:1; PPFD 252) | 110 days | Temperature: 24 °C Photoperiod: 16/8 h (day/night) | LED2 and LED5 effectively sustained C. sativa growth in plant height, stem diameter, and leaf numbers compared to CK. LED1, LED4, and LED3 treatments, which significantly reduced aboveground biomass, whereas LED2 and LED5 enhanced it. Flower biomass for LED2 and LED5 significantly increased compared to CK, while other LED treatments notably decreased flowers yield. | LED2 could significantly increase the CBD content of both leaves and flowers compared to CK. LED5 only notably increased the CBD content of leaves among the LED treatments. | [8] |
| 21. | C. sativa L. strain India | Green, far-red, and UV-A | 25 days | Temperature: 23–27 °C Photoperiod: 16 h PPFD: 300 μmol m−2 s−1 | -- | THC levels were higher in plants exposed to white, R8/B2, and R7/B2/G1 light. THCA accumulation was elevated in R6/B2/G1/FR1 and R5/B2/W2/FR1 light treatments, which coincided with reduced photosynthetic rate and increased ROS, total phenols, total flavonoids, DPPH radical scavenging capacity, and antioxidant enzymatic activities. The R6/B2/G1/FR1 light treatment resulted in higher CBDA accumulation, increased stress-modulated substances, and reduced physiological traits. | [62] |
| 22. | Stillwater cultivar | PPFDs ranging from 120 to 1800 μmol m−2 s−1 | 84 days | Day temperature: 25.3 ± 0.4 °C RH: 60.5 ± 4.8% Night temperature: 25.2 ± 0.3 °C RH: 53.1 ± 3.3% | Dry inflorescence yield exhibited a linear increase with increasing canopy-level PPFD up to 1800 μmol m−2 s−1, whereas leaf-level photosynthesis saturated well below 1800 μmol m−2 s−1. | There were minor LI treatment effects on the potency of cannabinoids and terpenes. The individual light effect is not specified. | [63] |
| 23. | C. sativa L. strain India | L1, natural light. L2, W; L3, R8/B2; L4, R7/B2/G1; L5, R7/B2/FR1; L6, R6/B2/G1/FR1; L7, R5/B2/W2/FR1; L8, R5/B2/G1/FR1/UV1; L9, R6/B2/FR1/UV1; L10, R4/B2/W2/FR1/UV1; L11, R2/B2/G2/W2/FR1/UV1 | 20 days | Temperature: 30 °C (day), 25 °C (night) RH: 60–70% Photoperiod: 12 h light | L10 and L11 exhibited more leaves and nodes, while L2, L3, and L5 showed increased leaf length and leaf width. Higher shoot lengths were noted in L3, L6, and L9. | L3 treatment exhibited higher levels of chlorophyll a, b, and photosynthetic quantum yield (Fv/Fm). L4, L6, L8, and L11 demonstrated a higher node count with elevated osmolyte content, including proline, ascorbic acid, total soluble carbohydrate, and sucrose. | [37] |
| 24. | Morphet Late, ECO-YP16, ECO-GH15, and ECO-MC16 | 600 W high-pressure sodium and eight 40 W globes suspended over the plants. Light intensity at canopy level (40 cm from the light source) was 300–350 μmol m−2 s−1 PAR | --- | Env.1: Daylength: 12.5 h Temperature: 27 °C/24 °C night (day/night) Env.2: Daylength: 11.5 h Temperature: 27 °C/24 °C night (day/night) Env.3: Daylength: 11.5 h Temperature: 22 °C/10 °C (day/night) | Tropical daylengths, temperatures, and nitrogen levels impacted C. sativa growth parameters. They play a crucial role in regulating flowering initiation, flower development, stalk diameter, and above-ground biomass in C. sativa varieties. | Environmental factors such as day length and temperature significantly influence cannabinoid concentrations. Selecting low-THC varieties was emphasized for production in tropical/subtropical environments. | [64] |
| 25. | King Harmony (Chemotype II, 1:1.5THC: CBD) | LED light, two low-white spectra (7B-20G-73R/narrow and 6B-19G-75R/2 peaks), and two high-white (15B-42G-43R/narrow and 17B-40G-43R/broad) spectra (600 and 1200 μmol m−2 s−1) | 56 days | Long-day phase: 28/24 °C, 27/22 °C, 26/22 °C, 25/22 °C Short-day phase: Days 0–28; 28/24 °C, Days 29–42; 27/22 °C, Days 49–56; 26/22 °C | The 6B/19G/75R (dual 640 and 660 nm peaks) enhanced inflorescence weight through increased dry matter production, while 7B-20G-73R (single 660 nm peak) reduced it. Two high-white spectra (15B-42G-43R/narrow and 17B-40G-43R/broad) maintained inflorescence weight and increased dry matter production at high PPFD. | The 6B-19G-75R (dual peaks: 640 and 660 nm) spectrum elicited an increase in terpenoid concentrations at high PPFD, whereas other light spectra did not induce significant alterations in cannabinoid levels. | [65] |