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Abstract Background Quantifying plant transpiration via thermal imaging is desirable for applications in agriculture, plant breeding, and plant science. However, thermal imaging under natural non-steady state conditions is currently limited by the difficulty of quantifying thermal properties of leaves, especially specific heat capacity (Cp). Existing literature offers only rough estimates of Cp and lacks simple and accurate methods to determine it. Results We developed a non-invasive method to quantify k (the product of leaf thickness (lt), leaf density(ρ), and Cp), by fitting a leaf energy balance model to a leaf temperature (Tleaf) transient during and after a ~ 10 s light pulse. Cp was then estimated by dividing k by lt*ρ. Using this method, we quantified Cp for 13 horticultural and tropical plant species, and explored the relationship between Cp and leaf water content, specific leaf area and Tleaf response rate during the light pulse. Values of Cp ranged between 3200–4000 J kg−1 K−1, and were positively correlated with leaf water content. In species with very thick leaves, such as Phalaenopsis amabilis, we found leaf thickness to be a major factor in the temperature response to a short light pulse. Conclusions Our method allows for easy determination of leaf Cp of different species, and may help pave the way to apply more accurate thermal imaging under natural non-steady state conditions.

Vertical farming is considered to be a key enabler for transforming agrifood systems, especially in or nearby urbanized areas. Vertical farming systems (VFS) are advanced indoor cropping systems that allow for highly intensified and standardized plant production. The close control of environmental parameters makes crop production stable and repeatable, ensuring year-round uniform product quality and quantity irrespective of location. However, due to continuous changes in plant physiology and development, as well as frequent changes in electricity prices, the optimum conditions for crop production and its associated costs can change within days or even minutes. This makes it beneficial to dynamically adjust setpoints for light (intensity, spectrum, pattern, and daylength), CO2, temperature, humidity, air flow, and water and nutrient availability. In this review, we highlight the beneficial effects that dynamic growth conditions can have on key plant processes, including improvements in photosynthetic gas exchange, transpiration, organ growth, development, light interception, flowering, and product quality. Our novel findings based on modeling and experimentation demonstrate that a dynamic daily light intensity pattern that responds to frequent changes in electricity prices can save costs without reducing biomass. Further, we argue that a smart, dynamic VFS climate management requires feedback mechanisms: several mobile and immobile sensors could work in combination to continuously monitor the crop, generating data that feeds into crop growth models, which, in turn, generate climate setpoints. In addition, we posit that breeding for the VFS environment is at a very early stage and highlight traits for breeding for this specialized environment. We envision a continuous feedback loop between dynamic crop management, crop monitoring, and trait selection for genotypes that are specialized for these conditions.

The cultivation of medical cannabis (Cannabis sativa L.) is expanding in controlled environments, driven by evolving governmental regulations for healthcare supply. Increasing inflorescence weight and plant specialized metabolite (PSM) concentrations is critical, alongside maintaining product consistency. Medical cannabis is grown under different spectra and photosynthetic photon flux densities (PPFD), the interaction between spectrum and PPFD on inflorescence weight and PSM attracts attention by both industrialists and scientists. Plants were grown in climate-controlled rooms without solar light, where four spectra were applied: two low-white spectra (7B-20G-73R/Narrow and 6B-19G-75R/2Peaks), and two high-white (15B-42G-43R/Narrow and 17B-40G-43R/Broad) spectra. The low-white spectra differed in red wavelength peaks (100% 660 nm, versus 50:50% of 640:660 nm), the high-white spectra differed in spectrum broadness. All four spectra were applied at 600 and 1200 μmol m-2 s-1. Irrespective of PPFD, white light with a dual red peak of 640 and 660 nm (6B-19G-75R/2Peaks) increased inflorescence weight, compared to white light with a single red peak of 660 nm (7B-20G-73R/Narrow) (tested at P = 0.1); this was associated with higher total plant dry matter production and a more open plant architecture, which likely enhanced light capture. At high PPFD, increasing white fraction and spectrum broadness (17B-40G-43R/Broad) produced similar inflorescence weights compared to white light with a dual red peak of 640 and 660 nm (6B-19G-75R/2Peaks). This was caused by an increase of both plant dry matter production and dry matter partitioning to the inflorescences. No spectrum or PPFD effects on cannabinoid concentrations were observed, although at high PPFD white light with a dual red peak of 640 and 660 nm (6B-19G-75R/2Peaks) increased terpenoid concentrations compared to the other spectra. At low PPFD, the combination of white light with 640 and 660 nm increased photosynthetic efficiency compared with white light with a single red peak of 660nm, indicating potential benefits in light use efficiency and promoting plant dry matter production. These results indicate that the interaction between spectrum and PPFD influences plant dry matter production. Dividing the light energy in the red waveband over both 640 and 660 nm equally shows potential in enhancing photosynthesis and plant dry matter production.

High planting densities achieve high light interception and harvestable yield per area but at the expense of product quality. This study aimed to maintain high light interception without negative impacts on fruit quality. Dwarf tomato was grown at four densities in a climate-controlled room—at two constant densities (high and low) and two dynamic spacing treatments (maintaining 90% and 75% ground coverage by decreasing planting density in 3–4 steps)—resulting in ~100, 19, 54, and 41 plants/m2 averaged over 100 days of cultivation, respectively. Constant high density resulted in the highest light use efficiency (LUE; 7.7 g fruit fresh weight per mol photons incident on the canopy) and the highest harvestable fruit yield (11.1 kg/m2) but the lowest fruit size and quality. Constant low density resulted in the lowest LUE and yield (2.3 g/mol and 3.2 kg/m2, respectively), but higher fruit size and quality than high density. Compared to low density, maintaining 90% ground coverage increased yield (9.1 kg/m2) and LUE (6.4 g/mol). Maintaining 75% ground coverage resulted in a 7.2 kg/m2 yield and 5.1 g/mol LUE. Both dynamic spacing treatments attained the same or slightly reduced fruit quality compared to low density. Total plant weight per m2 increased with planting density and saturated at a constant high density. Assimilate shortage at the plant level and flower abortion lowered harvestable fruit yield per plant, sweetness, and acidity under constant high density. Harvestable fruit yield per plant was the highest under dynamic spacing and low density. Under constant high density, morphological responses to lower light availability per plant—i.e., higher specific leaf area, internode elongation, and increased slenderness—coincided with the improved whole-plant LUE (g plant dry weight per mol photons). We conclude that a constant high planting density results in the highest harvestable fruit yield per area, but with reduced fruit quality. Dynamic spacing during cultivation produces the same fruit quality as constant low density, but with more than double the harvestable yield per area.

In controlled environment agriculture, customized light treatments using light-emitting diodes are crucial to improving crop yield and quality. Red (R; 600-700 nm) and blue light (B; 400-500 nm) are two major parts of photosynthetically active radiation (PAR), often preferred in crop production. Far-red radiation (FR; 700-800 nm), although not part of PAR, can also affect photosynthesis and can have profound effects on a range of morphological and physiological processes. However, interactions between different red and blue light ratios (R:B) and FR on promoting yield and nutritionally relevant compounds in crops remain unknown. Here, lettuce was grown at 200 µmol m-2 s-1 PAR under three different R:B ratios: R:B87.5:12.5 (12.5% blue), R:B75:25 (25% blue), and R:B60:40 (40% blue) without FR. Each treatment was also performed with supplementary FR (50 µmol m-2 s-1; R:B87.5:12.5+FR, R:B75:25+FR, and R:B60:40+FR). White light with and without FR (W and W+FR) were used as control treatments comprising of 72.5% red, 19% green, and 8.5% blue light. Decreasing the R:B ratio from R:B87.5:12.5 to R:B60:40, there was a decrease in fresh weight (20%) and carbohydrate concentration (48% reduction in both sugars and starch), whereas pigment concentrations (anthocyanins, chlorophyll, and carotenoids), phenolic compounds, and various minerals all increased. These results contrasted the effects of FR supplementation in the growth spectra; when supplementing FR to different R:B backgrounds, we found a significant increase in plant fresh weight, dry weight, total soluble sugars, and starch. Additionally, FR decreased concentrations of anthocyanins, phenolic compounds, and various minerals. Although blue light and FR effects appear to directly contrast, blue and FR light did not have interactive effects together when considering plant growth, morphology, and nutritional content. Therefore, the individual benefits of increased blue light fraction and supplementary FR radiation can be combined and used cooperatively to produce crops of desired quality: adding FR increases growth and carbohydrate concentration while increasing the blue fraction increases nutritional value.

Salinity is a current and growing problem, affecting crops worldwide by reducing yields and product quality. Plants have different mechanisms to adapt to salinity; some crops are highly studied, and their salinity tolerance mechanisms are widely known. However, there are other crops with commercial importance that still need characterization of their molecular mechanisms. Usually, transcription factors are in charge of the regulation of complex processes such as the response to salinity. MYB-TFs are a family of transcription factors that regulate various processes in plant development, and both central and specialized metabolism. MYB-TFs have been studied extensively as mediators of specialized metabolism, and some are master regulators. The influence of MYB-TFs on highly orchestrated mechanisms, such as salinity tolerance, is an attractive research target. The versatility of petunia as a model species has allowed for advances to be made in multiple fields: metabolomic pathways, quality traits, stress resistance, and signal transduction. It has the potential to be the link between horticultural crops and lab models, making it useful in translating discoveries related to the MYB-TF pathways into other crops. We present a phylogenetic tree made with Petunia axillaris and Petunia inflata R2R3-MYB subfamily sequences, which could be used to find functional conservation between different species. This work could set the foundations to improve salinity resistance in other commercial crops in later studies.

Artemisinin is a sesquiterpene lactone produced in glandular trichomes of Artemisia annua, and is extensively used in the treatment of malaria. Growth and secondary metabolism of A. annua are strongly regulated by environmental conditions, causing unstable supply and quality of raw materials from field grown plants. This study aimed to bring A. annua into greenhouse cultivation and to increase artemisinin production by manipulating greenhouse light environment using LEDs. A. annua plants were grown in a greenhouse compartment for five weeks in vegetative stage with either supplemental photosynthetically active radiation (PAR) (blue, green, red or white) or supplemental radiation outside PAR wavelength (far-red, UV-B or both). The colour of supplemental PAR hardly affected plant morphology and biomass, except that supplemental green decreased plant biomass by 15% (both fresh and dry mass) compared to supplemental white. Supplemental far-red increased final plant height by 23% whereas it decreased leaf area, plant fresh and dry weight by 30%, 17% and 7%, respectively, compared to the treatment without supplemental radiation. Supplemental UV-B decreased plant leaf area and dry weight (both by 7%). Interestingly, supplemental green and UV-B increased leaf glandular trichome density by 11% and 9%, respectively. However, concentrations of artemisinin, arteannuin B, dihydroartemisinic acid and artemisinic acid only exhibited marginal differences between the light treatments. There were no interactive effects of far-red and UV-B on plant biomass, morphology, trichome density and secondary metabolite concentrations. Our results illustrate the potential of applying light treatments in greenhouse production of A. annua to increase trichome density in vegetative stage. However, the trade-off between light effects on plant growth and trichome initiation needs to be considered. Moreover, the underlying mechanisms of light spectrum regulation on artemisinin biosynthesis need further clarification to enhance artemisinin yield in greenhouse production of A. annua.

Plant-to-plant variation (interplant variation) may play an important role in determining individual plant and whole canopy performance, where interplant variation in architecture and photosynthesis traits has direct effects on light absorption and photosynthesis. We aimed to quantify the importance of observed interplant variation on both whole-plant and canopy light absorption and photosynthesis. Plant architecture was measured in two experiments with fruiting tomato crops (Solanum lycopersicum) grown in glasshouses in the Netherlands, in week 16 (Exp. 1) or week 19 (Exp. 2) after transplanting. Experiment 1 included four cultivars grown under three supplementary lighting treatments, and Experiment 2 included two different row orientations. Measured interplant variations of the architectural traits, namely, internode length, leaf area, petiole angle, and leaflet angle, as well as literature data on the interplant variation of the photosynthesis traits alpha, Jmax28, and Vcmax28, were incorporated in a static functional–structural plant model (FSPM). The FSPM was used to analyze light absorption and net photosynthesis of whole plants in response to interplant variation in architectural and photosynthesis traits. Depending on the trait, introducing interplant variation in architecture and photosynthesis traits in a functional–structural plant model did not affect or negatively affected canopy light absorption and net photosynthesis compared with the reference model without interplant variation. Introducing interplant variation of architectural and photosynthesis traits in FSPM results in a more realistic simulation of variation of plants within a canopy. Furthermore, it can improve the accuracy of simulation of canopy light interception and photosynthesis although these effects at the canopy level are relatively small (

Abstract Vertical farming is a relatively new fresh fruit and vegetable production system, where lamps (mostly light emitting diodes [LED]) are the sole light source. A high light use efficiency (LUEinc), defined as shoot dry weight per incident photosynthetic photon flux density (PPFD; g mol−1) integral, is crucial for the economic viability of vertical farming. Very different values for LUEinc have been reported in the literature and it is not clear whether LUEinc is higher in vertical farming than in greenhouse or open field cultivation. Values of LUEinc of lettuce grown in a vertical farm (53 studies), greenhouse (13 studies) and open field (8 studies) were collected from literature, as well as relevant cultivation aspects such as lettuce weight at harvest, cultivation period (plant age at harvest), daily light integral, cumulative daily light integral for the whole cultivation period, planting density and CO2 concentration. The average LUEinc for lettuce grown in a vertical farm was 0.55 g mol−1 which was higher than 0.39 g mol−1 for greenhouse‐grown lettuce. Both were substantially higher than for field‐grown lettuce (0.23 g mol−1). The maximum measured LUEinc for lettuce grown in a vertical farm (1.63 g mol−1) is close to the published maximum theoretical value, which ranges from 1.26 to 1.81 g mol−1. Since all environmental factors can be fully controlled, vertical farming has the capability to achieve the theoretical maximum LUEinc. Using the highest reported LUEinc based on shoot fresh weight (44 g mol−1 at 200 μmol m−2 s−1 PPFD and 16 h photoperiod), it is estimated that each layer of a vertical farm can potentially produce annually up to 700 kg of lettuce per m2 at 500 μmol m−2 s−1 of continuous light.

Fluctuations in light intensity and temperature lead to periods of asynchrony between carbon (C) supply by photosynthesis and C demand by the plant organs. Storage and remobilization of non-structural carbohydrates (NSC) are important processes that allow plants to buffer these fluctuations. We aimed to test the hypothesis that C storage and remobilization can buffer the effects of temperature and light fluctuations on growth of tomato plants. Tomato plants were grown at temperature amplitudes of 3 or 10°C (deviation around the mean of 22°C) combined with integration periods (IP) of 2 or 10 days. Temperature and light were applied in Phase (high temperature simultaneously with high light intensity, (400 μmol m–2 s–1), low temperature simultaneously with low light intensity (200 μmol m–2 s–1) or in Antiphase (high temperature with low light intensity, low temperature with high light intensity). A control treatment with constant temperature (22°C) and a constant light intensity (300 μmol m–2 s–1) was also applied. After 20 days all treatments had received the same temperature and light integral. Differences in final structural dry weight were relatively small, while NSC concentrations were highly dynamic and followed changes of light and temperature (a positive correlation with decreasing temperature and increasing light intensity). High temperature and low light intensity lead to depletion of the NSC pool, but NSC level never dropped below 8% of the plant weight and this fraction was not mobilizable. Our results suggest that growing plants under fluctuating conditions do not necessarily have detrimental effects on plant growth and may improve biomass production in plants. These findings highlight the importance in the NSC pool dynamics to buffer fluctuations of light and temperature on plant structural growth.

Abstract Background There is little information on the effect of nutrient solutions composition on Arabidopsis growth. Therefore, we compared growth performance of Arabidopsis thaliana (Col-0) grown on the most commonly used nutrient solutions in deep water culture: Hoagland and Arnon, Murashige and Skoog, Tocquin, Hermans, and Conn. In addition to these nutrient solution composition experiments, we established Arabidopsis growth response curves for nutrient solution concentration and salt stress (NaCl). Results Arabidopsis rosette fresh and dry weight showed an approximate linear decline with NaCl dose in deep water culture, i.e. 9% reduction relative to control per unit of electrical conductivity (EC in dS m−1, for scale comprehension 1 dS m−1 equals ~ 10 mM NaCl). The Tocquin, ½Hoagland and Conn nutrient solutions had equal and optimal growth performance. Optimal nutrient solution concentration for Tocquin and Hoagland was 0.8 to 0.9 dS m−1. Close to the EC of ½Hoagland (1.1 dS m−1), which is frequently used in Arabidopsis research. Conn solution showed optimal growth at much higher EC (2 dS m−1) indicating that it is a balanced nutrient solution that matches the needs of Arabidopsis. Full Murashige and Skoog solution (5.9 dS m−1) was lethal and diluted solutions (EC of 1.6 and 1.1 dS m−1) caused stress symptoms and severe growth retardation at later developmental stages. Conclusions Arabidopsis thaliana (Col-0) plants grown in deep water culture showed a sixfold growth difference when commonly used nutrient solutions were compared. Murashige and Skoog solution should not be used as nutrient solution in deep water culture. Conn, Tocquin and ½Hoagland are balanced nutrient solutions which result in optimal Arabidopsis growth in hydroponic systems.

Blue light, measuring from 400 to 500 nm, is generally assumed to increase the content of antioxidants in plants independent of the species. Blue light stimulates the biosynthesis of phenolic compounds such as flavonoids and their subclass anthocyanins from the phenylpropanoid pathway. Flavonoids, anthocyanins, and phenolic acids are strong reactive oxygen species (ROS) scavengers and may lessen the symptoms of abiotic stresses such as chilling. We tested the hypothesis that a high percentage of blue light induces the accumulation of antioxidants and that this effect depends on the photosynthetic photon flux density (PPFD, 400–700 nm). The effect may be more pronounced at a lower PPFD. We investigated the changes in primary and secondary metabolites of basil in response to the percentage of blue light (9, 33, 65, and 100%) applied either as a 5-day End-Of-Production (EOP) treatment or continuous throughout the growth cycle in the green cv. Dolly. We also studied if the response to the percentage of blue light (9 or 90%) was dependent on the total PPFD (100 or 300 μmol m–2 s–1 PPFD) when applied as a 5-day EOP treatment in the green cv. Dolly and the purple cv. Rosie. For both green and purple basil, it was found that the percentage of blue light had little effect on the levels of antioxidants (rosmarinic acid, total ascorbic acid, total flavonoids, and total anthocyanins) at harvest and no interactive effect with PPFD was found. Antioxidants generally decreased during postharvest storage, wherein the decrease was more pronounced at 4 than at 12°C. Chilling injury, as judged from a decrease in Fv/Fm values and from the occurrence of black necrotic areas, was not affected by the percentage of blue light. Particularly, chilling tolerance in the purple cultivar was increased in plants grown under higher PPFD. This may be related to the increased levels of soluble sugar and starch in leaves from high PPFD treated plants.

Under natural conditions, irradiance frequently fluctuates, causing net photosynthesis rate (A) to respond slowly and reducing the yields. We quantified the genotypic variation of photosynthetic induction in 19 genotypes among the following six horticultural crops: basil, chrysanthemum, cucumber, lettuce, tomato, and rose. Kinetics of photosynthetic induction and the stomatal opening were measured by exposing shade-adapted leaves (50 μmol m–2 s–1) to a high irradiance (1000 μmol m–2 s–1) until A reached a steady state. Rubisco activation rate was estimated by the kinetics of carboxylation capacity, which was quantified using dynamic A vs. [CO2] curves. Generally, variations in photosynthetic induction kinetics were larger between crops and smaller between cultivars of the same crop. Time until reaching 20–90% of full A induction varied by 40–60% across genotypes, and this was driven by a variation in the stomatal opening rather than Rubisco activation kinetics. Stomatal conductance kinetics were partly determined by differences in the stomatal size and density; species with densely packed, smaller stomata (e.g., cucumber) tended to open their stomata faster, adapting stomatal conductance more rapidly and efficiently than species with larger but fewer stomata (e.g., chrysanthemum). We conclude that manipulating stomatal traits may speed up photosynthetic induction and growth of horticultural crops under natural irradiance fluctuations.

Shorter photoperiod and lower daily light integral (DLI) limit the winter greenhouse production. Extending the photoperiod by supplemental light increases biomass production but inhibits flowering in short-day plants such as Chrysanthemum morifolium. Previously, we reported that flowering in growth-chamber grown chrysanthemum with red (R) and blue (B) LED-light could also be induced in long photoperiods by applying only blue light during the last 4h of 15h long-days. This study investigates the possibility to induce flowering by extending short-days in greenhouses with 4h of blue light. Furthermore, flower induction after 4h of red light extension was tested after short-days RB-LED light in a growth-chamber and after natural solar light in a greenhouse. Plants were grown at 11h of sole source RB light (60:40) in a growth-chamber or solar light in the greenhouse (short-days). Additionally, plants were grown under long-days, which either consisted of short-days as described above extended with 4h of B or R light to long-days or of 15h continuous RB light or natural solar light. Flower initiation and normal capitulum development occurred in the blue-extended long-days in the growth-chamber after 11h of sole source RB, similarly as in short-days. However, when the blue extension was applied after 11h of full-spectrum solar light in a greenhouse, no flower initiation occurred. With red-extended long-days after 11h RB (growth-chamber) flower initiation occurred, but capitulum development was hindered. No flower initiation occurred in red-extended long-days in the greenhouse. These results indicate that multiple components of the daylight spectrum influence different phases in photoperiodic flowering in chrysanthemum in a time-dependent manner. This research shows that smart use of LED-light can open avenues for a more efficient year-round cultivation of chrysanthemum by circumventing the short-day requirement for flowering when applied in emerging vertical farm or plant factories that operate without natural solar light. In current year-round greenhouses’ production, however, extension of the natural solar light during the first 11 h of the photoperiod with either red or blue sole LED light, did inhibit flowering.

The recent development of light-emitting diodes (LEDs) and their application in modern horticulture stimulated studies demonstrating that additional far-red (FR) radiation (700–800 nm) increases plant dry mass. This effect of FR has been explained by improved photosynthesis and/or plant architecture. However, the genotypic variation in this response is largely unknown. Here, we aim to explore and explain the genotypic variation in growth responses to additional FR. We expected the genotypic variation in the responses of plant dry mass to additional FR. Further, we hypothesized that a significant improvement of both net assimilation rate (NAR) and leaf area ratio (LAR) is responsible for a strong dry mass increase under additional FR, while some genotypes respond only marginally or even negatively in NAR or LAR under FR, thus resulting in a weak FR effect on plant dry mass. To test these hypotheses, we grew 33 different tomato genotypes for 21 days with 0, 25, or 100 μmol m–2 s–1 of FR added to a common white + red LED background lighting of 150 μmol m–2 s–1. Genotypes responded similarly with respect to plant height, stem dry mass, and shoot:root ratio; i.e., they all increased with increasing FR. However, the response of total plant dry mass varied among genotypes. We categorized the genotypes into three groups (strongly, moderately, and weakly responding groups) based on their relative response in total plant dry mass to FR. Growth component analysis revealed that the strongly responding genotypes increased strongly in NAR rather than LAR. The weakly responding genotypes, however, showed a substantial increase in LAR but not NAR. The increase in LAR was due to the increase in specific leaf area. Leaf mass fraction, which is the other component of LAR, decreased with FR and did not differ between groups. In conclusion, tomato genotypes that increased strongly in NAR in response to FR were able to achieve a more substantial increase in dry mass than did other genotypes. This is the first study to explain the differences in growth responses of a large number of tomato genotypes toward FR in their light environment.

The effect of light intensity applied shortly before harvest on the nutritional quality, postharvest performance, and shelf life of loose-leaf lettuce (Lactuca sativa L. cv. Expertise RZ Salanova®) was investigated. Lettuce was grown either in a greenhouse with supplemental high-pressure sodium light (Experiment 1, EXP 1) or in a climate room under white LED light (Experiment 2, EXP 2). In both experiments full grown plants were transferred to a climate room for the End of Production (EoP) light treatments during the last week of cultivation. During EoP lighting plants were exposed to different intensities (0, 110, and 270 μmol m–2 s–1 in EXP 1; 50, 210, and 470 μmol m–2 s–1 in EXP 2) from white-red LEDs for 6 (EXP 2) or 7 days (EXP 1). Mature leaves were then harvested and stored in darkness at 10°C to study the postharvest performance. Changes in dry matter content, total ascorbic acid, and carbohydrates (including glucose, fructose sucrose, and starch) levels were determined during EoP lighting and during the subsequent shelf life as indicators of lettuce nutritional quality. Quality aspects (appearance, texture, and odor) were accessed during the shelf life as indicators of postharvest performance. In both experiments, high light intensities applied in EoP lighting increased dry matter percentage and contents of ascorbic acid (AsA) and carbohydrates at harvest and these increased levels were maintained during the shelf life. Increased light intensity in EoP treatment also extended the shelf life. The levels of AsA and carbohydrates at harvest correlated positively with the subsequent shelf life, indicating that the prolonged shelf life relies on the improved energy and antioxidant status of the crop at harvest.

The economic viability and energy use of vertical farms strongly depend on the efficiency of the use of light. Increasing far-red radiation (FR, 700–800 nm) relative to photosynthetically active radiation (PAR, 400–700 nm) may induce shade avoidance responses including stem elongation and leaf expansion, which would benefit light interception, and FR might even be photosynthetically active when used in combination with PAR. The aims of this study are to investigate the interaction between FR and planting density and to quantify the underlying components of the FR effects on growth. Lettuce (Lactuca sativa cv. Expertise RZ) was grown in a climate chamber under two FR treatments (0 or 52 μmol m–2 s–1) and three planting densities (23, 37, and 51 plants m–2). PAR of 89% red and 11% blue was kept at 218 μmol m–2 s–1. Adding FR increased plant dry weight after 4 weeks by 46–77% (largest effect at lowest planting density) and leaf area by 58–75% (largest effect at middle planting density). Radiation use efficiency (RUE: plant dry weight per unit of incident radiation, 400–800 nm) increased by 17–42% and incident light use efficiency (LUEinc: plant dry weight per unit of incident PAR, 400–700 nm) increased by 46–77% by adding FR; the largest FR effects were observed at the lowest planting density. Intercepted light use efficiency (LUEint: plant dry weight per unit of intercepted PAR) increased by adding FR (8–23%). Neither specific leaf area nor net leaf photosynthetic rate was influenced by FR. We conclude that supplemental FR increased plant biomass production mainly by faster leaf area expansion, which increased light interception. The effects of FR on plant dry weight are stronger at low than at high planting density. Additionally, an increased LUEint may contribute to the increased biomass production.

Vertical farming is becoming increasingly popular for production of leafy vegetables and herbs, with basil (Ocimum basilicum L.) as one of the most popular herbs. In basil most research has focused on increasing secondary metabolites with light spectra. However, knowledge about the effect of light intensity (photosynthetic photon flux density, PPFD) and spectra on growth and morphology is key for optimizing quality at harvest. The impact of PPFD and spectrum on plant growth and development is species dependent and currently few studies in basil are available. Understanding the response to End-Of-Production (EOP) light of growth and morphology is important for successful vertical farming. We performed a comprehensive series of experiments, where the effects of EOP PPFD, fraction of blue and their interaction on the growth and morphology were analyzed in two green and one purple basil cultivar. In addition, the impact of different EOP intensities and duration of far-red were investigated. We found that increasing the PPFD increased fresh mass, dry matter content and plant height in all three cultivars. The responses were linear or quadratic depending on the cultivar. A high fraction of blue (>90%) increased plant height and decreased the dry mass partitioning to the leaves. The only interaction found between the fraction of blue and overall PPFD was on plant height in the green cultivar whereas other growth parameters and morphology responded stronger to PPFD than to the fraction of blue light. Plant dry matter production was increased with the addition of far-red. Far-red EOP intensity treatments enhanced the fraction of dry mass partitioned to the leaves, whereas a prolonged far-red treatment enhanced partitioning to the stem. Both plant fresh mass and dry matter content were improved by applying high PPFD shortly before harvest. Light spectra were found to be of less importance than PPFD with respect to plant dry matter content. Light use efficiency (LUE) based on fresh mass decreased with increasing PPFD whereas LUE based on dry mass increased with increasing PPFD, when given as EOP treatments. The overall physiological mechanisms of the light intensity and spectral effects are discussed.

Due to their slow movement and closure upon shade, partially closed stomata can be a substantial limitation to photosynthesis in variable light intensities. The abscisic acid deficient flacca mutant in tomato (Solanum lycopersicum) displays very high stomatal conductance (gs). We aimed to determine to what extent this substantially increased gs affects the rate of photosynthetic induction. Steady-state and dynamic photosynthesis characteristics were measured in flacca and wildtype leaves, by the use of simultaneous gas exchange and chlorophyll fluorometry. The steady-state response of photosynthesis to CO2, maximum quantum efficiency of photosystem II photochemistry (Fv/Fm), as well as mesophyll conductance to CO2 diffusion were not significantly different between genotypes, suggesting similar photosynthetic biochemistry, photoprotective capacity, and internal CO2 permeability. When leaves adapted to shade (50 µmol m−2 s−1) at 400 µbar CO2 partial pressure and high humidity (7 mbar leaf-to-air vapour pressure deficit, VPD) were exposed to high irradiance (1500 µmol m−2 s−1), photosynthetic induction was faster in flacca compared to wildtype leaves, and this was attributable to high initial gs in flacca (~0.6 mol m−2 s−1): in flacca, the times to reach 50 (t50) and 90% (t90) of full photosynthetic induction were 91 and 46% of wildtype values, respectively. Low humidity (15 mbar VPD) reduced gs and slowed down photosynthetic induction in the wildtype, while no change was observed in flacca; under low humidity, t50 was 63% and t90 was 36% of wildtype levels in flacca. Photosynthetic induction in low CO2 partial pressure (200 µbar) increased gs in the wildtype (but not in flacca), and revealed no differences in the rate of photosynthetic induction between genotypes. Effects of higher gs in flacca were also visible in transients of photosystem II operating efficiency and non-photochemical quenching. Our results show that at ambient CO2 partial pressure, wildtype gs is a substantial limitation to the rate of photosynthetic induction, which flacca overcomes by keeping its stomata open at all times, and it does so at the cost of reduced water use efficiency.

Green light (G) could trigger a “shade avoidance syndrome” (SAS) similarly to far-red light. We aimed to test the hypothesis that G interacts with far-red light to induce SAS, with this interaction mediated by phytochromes (phys). The tomato (Solanum lycopersicum cv. Moneymaker) wild-type (WT) and phyA, phyB1B2, and phyAB1B2 mutants were grown in a climate room with or without 30 µmol m−2 s−1 G on red/blue and red/blue/far-red backgrounds, maintaining the same photosynthetically active radiation (400–700 nm) of 150 µmol m−2 s−1 and red/blue ratio of 3. G hardly affected the dry mass accumulation or leaf area of WT, phyA, and phyB1B2 with or without far-red light. A lower phytochrome photostationary state (PSS) by adding far-red light significantly increased the total dry mass by enhancing the leaf area in WT plants but not in phy mutants. When the background light did not contain far-red light, partially replacing red/blue with G did not significantly affect stem elongation. However, when the background light contained far-red light, partially replacing red/blue with G enhanced elongation only when associated with a decrease in PSS, indicating that G interacts with far-red light on elongation only when the PSS changes.

Indoor plant cultivation can result in significantly improved resource use efficiency (surface, water, and nutrients) as compared to traditional growing systems, but illumination costs are still high. LEDs (light emitting diodes) are gaining attention for indoor cultivation because of their ability to provide light of different spectra. In the light spectrum, red and blue regions are often considered the major plants’ energy sources for photosynthetic CO2 assimilation. This study aims at identifying the role played by red:blue (R:B) ratio on the resource use efficiency of indoor basil cultivation, linking the physiological response to light to changes in yield and nutritional properties. Basil plants were cultivated in growth chambers under five LED light regimens characterized by different R:B ratios ranging from 0.5 to 4 (respectively, RB0.5, RB1, RB2, RB3, and RB4), using fluorescent lamps as control (CK1). A photosynthetic photon flux density of 215 μmol m−2 s−1 was provided for 16 h per day. The greatest biomass production was associated with LED lighting as compared with fluorescent lamp. Despite a reduction in both stomatal conductance and PSII quantum efficiency, adoption of RB3 resulted in higher yield and chlorophyll content, leading to improved use efficiency for water and energy. Antioxidant activity followed a spectral-response function, with optimum associated with RB3. A low RB ratio (0.5) reduced the relative content of several volatiles, as compared to CK1 and RB ≥ 2. Moreover, mineral leaf concentration (g g−1 DW) and total content in plant (g plant−1) were influences by light quality, resulting in greater N, P, K, Ca, Mg, and Fe accumulation in plants cultivated with RB3. Contrarily, nutrient use efficiency was increased in RB ≤ 1. From this study it can be concluded that a RB ratio of 3 provides optimal growing conditions for indoor cultivation of basil, fostering improved performances in terms of growth, physiological and metabolic functions, and resources use efficiency.

Shading by sunlit leaves causes a low red (R) to far-red (FR) ratio that results in a low phytochrome stationary state (PSS). A low PSS induces an array of shade avoidance responses that influence plant architecture and development. It has often been suggested that this architectural response is advantageous for plant growth due to its positive effect on light interception. In contrast to sunlight, artificial light sources such as LEDs often lack FR, resulting in a PSS value higher than solar light (∼0.70). The aim of this study was to investigate how PSS values higher than solar radiation influence the growth and development of tomato plants. Additionally, we investigated whether a short period of FR at the end of the day (EOD-FR) could counteract any potentially negative effects caused by a lack of FR during the day. Tomato plants were grown at four PSS levels (0.70, 0.73, 0.80, and 0.88), or with a 15-min end-of-day far-red (EOD-FR) application (PSS 0.10). Photosynthetic Active Radiation (PAR; 150 μmol m-2 s-1) was supplied using red and blue (95/5%) LEDs. In an additional experiment, the same treatments were applied to plants receiving supplementary low-intensity solar light. Increasing PSS above solar PSS resulted in increased plant height. Leaf area and plant dry mass were lower in the treatments completely lacking FR than treatments with FR. EOD-FR-treated plants responded almost similarly to plants grown without FR, except for plant height, which was increased. Simulations with a 3D-model for light absorption revealed that the increase in dry mass was mainly related to an increase in light absorption due to a higher total leaf area. Increased petiole angle and internode length had a negative influence on total light absorption. Additionally, the treatments without FR and the EOD-FR showed strongly reduced fruit production due to reduced fruit growth associated with reduced source strength and delayed flowering. We conclude that growing tomato plants under artificial light without FR during the light period causes a range of inverse shade avoidance responses, which result in reduced plant source strength and reduced fruit production, which cannot be compensated by a simple EOD-FR treatment.

Metabolic profiles of tomatoes change during ripening and light can modulate the activity of relevant biochemical pathways. We investigated the effects of light directly supplied to the fruits on the metabolome of the fruit pericarp during ripening. Mature green tomatoes were exposed to well-controlled conditions with light as the only varying factor; control fruits were kept in darkness. In experiment 1 the fruits were exposed to either white light or darkness for 15 days. In experiment 2, fruits were exposed to different light spectra (blue, green, red, far-red, white) added to white background light for seven days. Changes in the global metabolome of the fruit pericarp were monitored using LCMS and GCMS (554 compounds in total). Health-beneficial compounds (carotenoids, flavonoids, tocopherols and phenolic acids) accumulated faster under white light compared to darkness, while alkaloids and chlorophylls decreased faster. Light also changed the levels of taste-related metabolites including glutamate and malate. The light spectrum treatments indicated that the addition of blue light was the most effective treatment in altering the fruit metabolome. We conclude that light during ripening of tomatoes can have various effects on the metabolome and may help with shaping the levels of key compounds involved in various fruit quality characteristics.

Greenhouse crop production in northern countries often relies heavily on supplemental lighting for year-round yield and product quality. Among the different spectra used in supplemental lighting, red is often considered the most efficient, but plants do not develop normally when grown solely under monochromatic red light (“red light syndrome”). Addition of blue light has been shown to aid normal development, and typical lighting spectra in greenhouse production include a mixture of red and blue light. However, it is unclear whether sunlight, as part of the light available to plants in the greenhouse, may be sufficient as a source of blue light. In a greenhouse high-wire tomato (Solanum lycopersicum), we varied the percentage of blue supplemental light (in a red background) as 0, 6, 12, and 24%, while keeping total photosynthetically active radiation constant. Light was supplied as a mixture of overhead (99 μmol m-2 s-1) and intracanopy (48 μmol m-2 s-1) LEDs, together with sunlight. Averaged over the whole experiment (111 days), sunlight comprised 58% of total light incident onto the crop. Total biomass, yield and number of fruits increased with the addition of blue light to an optimum, suggesting that both low (0%) and high (24%) blue light intensities were suboptimal for growth. Stem and internode lengths, as well as leaf area, decreased with increases in blue light percentage. While photosynthetic capacity increased linearly with increases in blue light percentage, photosynthesis in the low blue light treatment (0%) was not low enough to suggest the occurrence of the red light syndrome. Decreased biomass at low (0%) blue light was likely caused by decreased photosynthetic light use efficiency. Conversely, decreased biomass at high (24%) blue light was likely caused by reductions in canopy light interception. We conclude that while it is not strictly necessary to add blue light to greenhouse supplemental red light to obtain a functional crop, adding some (6–12%) blue light is advantageous for growth and yield while adding 24% blue light is suboptimal for growth.

Anthocyanins are a group of polyphenolic pigments that are ubiquitously found in the plant kingdom. In plants, anthocyanins play a role not only in reproduction, by attracting pollinators and seed dispersers, but also in protection against various abiotic and biotic stresses. There is accumulating evidence that anthocyanins have health-promoting properties, which makes anthocyanin metabolism an interesting target for breeders and researchers. In this review, the state of the art knowledge concerning anthocyanins in the Solanaceous vegetables, i.e., pepper, tomato, eggplant, and potato, is discussed, including biochemistry and biological function of anthocyanins, as well as their genetic and environmental regulation. Anthocyanin accumulation is determined by the balance between biosynthesis and degradation. Although the anthocyanin biosynthetic pathway has been well-studied in Solanaceous vegetables, more research is needed on the inhibition of biosynthesis and, in particular, the anthocyanin degradation mechanisms if we want to control anthocyanin content of Solanaceous vegetables. In addition, anthocyanin metabolism is distinctly affected by environmental conditions, but the molecular regulation of these effects is poorly understood. Existing knowledge is summarized and current gaps in our understanding are highlighted and discussed, to create opportunities for the development of anthocyanin-rich crops through breeding and environmental management.

L-ascorbate (ASC) is essential for human health. Therefore, there is interest in increasing the ASC content of crops like tomato. High irradiance induces accumulation of ASC in green tomato fruits. The D-mannose/L-galactose biosynthetic pathway accounts for the most ASC in plants. The myo-inositol and galacturonate pathways have been proposed to exist but never identified in plants. The D-mannose/L-galactose starts from D-glucose. In a series of experiments, we tested the hypothesis that ASC levels depend on soluble carbohydrate content when tomato fruits ripen under irradiances that stimulate ASC biosynthesis. We show that ASC levels considerably increased when fruits ripened under light, but carbohydrate levels did not show a parallel increase. When carbohydrate levels in fruits were altered by flower pruning, no effects on ASC levels were observed at harvest or after ripening under irradiances that induce ASC accumulation. Artificial feeding of trusses with sucrose increased carbohydrate levels, but did not affect the light-induced ASC levels. We conclude that light-induced accumulation of ASC is independent of the carbohydrate content in tomato fruits. In tomato fruit treated with light, the increase in ASC was preceded by a concomitant increase in myo-inositol.