The role of red and white light in optimizing growth and accumulation of plant specialized metabolites at two light intensities in medical cannabis (Cannabis sativa L.).

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.

Green light is similarly effective in promoting plant biomass as red/blue light - a meta-analysis.

Whether green light promotes or represses plant growth is an unresolved but important question, warranting a global meta-analysis of published data. We collected 136 datasets from 48 publications on 17 crop species, and calculated the green light effect for a range of plant traits. For each trait the effect was calculated as the ratio between the trait value attained under a red/blue background light plus green, divided by the value attained under the background light only, both having the same light intensity. Generally, green light strongly increased intrinsic water use efficiency (15%), the shoot-to-root ratio (13%), and decreased stomatal conductance (-15%). Moreover, green light increased fresh weight to a small extent (4%), but not plant dry weight, resulting in a reduced dry matter content (-2%). Hence, green light is similarly effective at increasing biomass as red and blue light. Green light also showed to increase leaf area (7%) and specific leaf area (4%; i.e., thinner leaves). Furthermore, effects of green light were species-dependent, with positive effects on biomass for lettuce and microgreens, and negative effects in basil and tomato. Our data suggest that future research should focus on the role of green light in modulating water loss, its putative role as a shade signal, and the causes for its species-specific effects on crop biomass.

Dynamic plant spacing in tomato results in high yields while mitigating the reduction in fruit quality associated with high planting densities.

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.

Ménage à trois: light, terpenoids, and quality of plants.

In controlled environment agriculture (CEA), light is used to impact terpenoid production and improve plant quality. In this review we discuss various aspects of light as important regulators of terpenoid production in different plant organs. Spectral quality primarily modifies terpenoid profiles, while intensity and photoperiod influence abundances. The central regulator of light signal transduction elongated hypocotyl 5 (HY5) controls transcriptional regulation of terpenoids under UV, red (R), and blue (B) light. The larger the fraction of R and green (G) light, the more beneficial the effect on monoterpenoid and sesquiterpenoid biosynthesis, and such an effect may depend on the presence of B light. A large fraction of R light is mostly detrimental to tetraterpenoid production. We conclude that light is a promising tool to steer terpenoid production and potentially tailor the quality of plants.

Far-red light-enhanced apical dominance stimulates flower and fruit abortion in sweet pepper.

Far-red radiation affects many plant processes, including reproductive organ abortion. Our research aimed to determine the role of apical dominance in far-red light-induced flower and fruit abortion in sweet pepper (Capsicum annuum L.). We conducted several climate room experiments where plants were grown under white- or red-rich LED light, with or without additional far-red light. Additional far-red light enhanced apical dominance: it increased auxin levels in the apices of dominant shoots, and caused a greater difference in internode length and apical auxin levels between dominant and subordinate shoots. Additional far-red light stimulated fruit abortion in intact plants but not in decapitated plants, suggesting a crucial role of shoot apices in this effect. However, reducing basipetal auxin transport in the stems with N-1-naphthylphthalamic acid did not influence far-red light-stimulated fruit abortion, although auxin levels in the stem were largely reduced. Applying the synthetic auxin 1-naphthaleneacetic acid on decapitated apices did not influence fruit abortion. However, applying the auxin biosynthesis inhibitor yucasin to shoot apices reduced fruit abortion regardless of the light conditions, accompanied by slight shoot growth retardation. These findings suggest that the basipetal auxin stream does not mediate far-red light-stimulated fruit abortion. Far-red light-stimulated fruit abortion was associated with reduced sucrose accumulation and lower invertase activities in flowers. We suggest that under additional far-red light conditions, increased auxin levels in shoot apices promote fruit abortion probably through enhanced competition for assimilates between apices and flowers, which limits assimilate import into flowers.

Young Tomato Plants Respond Differently under Single or Combined Mild Nitrogen and Water Deficit: An Insight into Morphophysiological Responses and Primary Metabolism.

This study aimed to understand the morphophysiological responses and primary metabolism of tomato seedlings subjected to mild levels of nitrogen and/or water deficit (50% N and/or 50% W). After 16 days of exposure, plants grown under the combined deficit showed similar behavior to the one found upon exposure to single N deficit. Both N deficit treatments resulted in a significantly lower dry weight, leaf area, chlorophyll content, and N accumulation but in a higher N use efficiency when compared to control (CTR) plants. Moreover, concerning plant metabolism, at the shoot level, these two treatments also responded in a similar way, inducing higher C/N ratio, nitrate reductase (NR) and glutamine synthetase (GS) activity, expression of RuBisCO encoding genes as well as a downregulation of GS2.1 and GS2.2 transcripts. Interestingly, plant metabolic responses at the root level did not follow the same pattern, with plants under combined deficit behaving similarly to W deficit plants, resulting in enhanced nitrate and proline concentrations, NR activity, and an upregulation of GS1 and NR genes than in CTR plants. Overall, our data suggest that the N remobilization and osmoregulation strategies play a relevant role in plant acclimation to these abiotic stresses and highlight the complexity of plant responses under a combined N+W deficit.

Enzymatic and Non-Enzymatic Antioxidant Responses of Young Tomato Plants (cv. Micro-Tom) to Single and Combined Mild Nitrogen and Water Deficit: Not the Sum of the Parts.

This study aims to perform a broad analysis of the antioxidant (AOX) responses of young tomato plants exposed to single and combined mild nitrogen (N) and water deficits through the evaluation of oxidative biomarkers, non-enzymatic and enzymatic AOX components. 'Micro-Tom' seedlings were subjected to four treatments: control (CTR; 100%N + 100%W), N deficit (N; 50%N), water deficit (W; 50%W), and combined deficits (N + W; 50%N + 50%W). An enhancement of several non-enzymatic and enzymatic components was found in plants subjected to N + W deficit, which presented higher anthocyanins accumulation (up to 103%) as well as higher levels of superoxide dismutase (SOD) transcripts at root level and of ascorbate peroxidase (APX) and catalase (CAT) transcripts at shoot level. This increase in the gene expression was also translated in augmented SOD (up to 202%), APX (up to 155%) and CAT (up to 108%) activity compared to CTR plants and the single deficits. Overall, tomato plants were able to employ defense strategies to cope with this combined deficit, as demonstrated by the higher total AOX capacity (up to 87%) compared to the single deficits, which contributed to the maintenance of their redox homeostasis, with unchanged values of lipid peroxidation and hydrogen peroxide compared with CTR plants.

Consequences of interplant trait variation for canopy light absorption and photosynthesis.

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, J max28, and V cmax28, 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 (<4% for light absorption and<7% for net photosynthesis).

Machine learning versus crop growth models: an ally, not a rival.

The rapid increases of the global population and climate change pose major challenges to a sustainable production of food to meet consumer demands. Process-based models (PBMs) have long been used in agricultural crop production for predicting yield and understanding the environmental regulation of plant physiological processes and its consequences for crop growth and development. In recent years, with the increasing use of sensor and communication technologies for data acquisition in agriculture, machine learning (ML) has become a popular tool in yield prediction (especially on a large scale) and phenotyping. Both PBMs and ML are frequently used in studies on major challenges in crop production and each has its own advantages and drawbacks. We propose to combine PBMs and ML given their intrinsic complementarity, to develop knowledge- and data-driven modelling (KDDM) with high prediction accuracy as well as good interpretability. Parallel, serial and modular structures are three main modes can be adopted to develop KDDM for agricultural applications. The KDDM approach helps to simplify model parameterization by making use of sensor data and improves the accuracy of yield prediction. Furthermore, the KDDM approach has great potential to expand the boundary of current crop models to allow upscaling towards a farm, regional or global level and downscaling to the gene-to-cell level. The KDDM approach is a promising way of combining simulation models in agriculture with the fast developments in data science while mechanisms of many genetic and physiological processes are still under investigation, especially at the nexus of increasing food production, mitigating climate change and achieving sustainability.

Non-structural carbohydrate dynamics and growth in tomato plants grown at fluctuating light and temperature.

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.

Effects of Green Light on Elongation Do Not Interact with Far-Red, Unless the Phytochrome Photostationary State (PSS) Changes in Tomato.

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.

LED Intercanopy Lighting in Blackberry During Spring Improves Yield as a Result of Increased Number of Fruiting Laterals and Has a Positive Carryover Effect on Autumn Yield.

High market price and low availability of local winter and spring production has stimulated production of blackberries in glasshouses at northern latitudes. For this production, light is the main limiting factor. We investigated the potential of intercanopy lighting (ICL) using light emitting diodes (LEDs) to improve blackberry fruit yield in a crop with a spring and an autumn production cycle. During the spring production cycle three light treatments were applied: only natural light (no ICL), 93 or 185 µmol m-2 s-1 ICL In summer the lateral shoots were cut back and 93 µmol m-2 s-1 ICL was applied to all plants after cutting back, investigating a possible carryover effect of supplemental light in spring on autumn production. Fresh fruit yield in spring increased by 79 and 122% with 93 and 185 µmol m-2 s-1 ICL, respectively, compared to no ICL. This represents 3.6 and 2.8% increase in harvestable product for every additional 1% of light. A yield component analysis and leaf photosynthesis measurements were conducted. Maximum photosynthetic capacity (A max) for leaves at 185 µmol m-2 s-1 ICL was about 50% higher, and LAI was 41% higher compared to no ICL. ICL increased the number of fruiting laterals per cane, and this explained 75% of the increase in yield. ICL at 185 µmol m-2 s-1 resulted in a higher yield compared to no ICL, primarily as a result of higher total dry matter production. Furthermore, a higher fraction of dry matter partitioned to the fruits (0.59 compared to 0.52) contributed to yield increase, whereas fruit dry matter content and fruit quality (sugar and acid content) was not affected by ICL. Averaged over the three light treatments autumn yield was 47% lower than spring yield. Autumn yield was 10% higher for plants at ICL 93 µmol m-2 s-1 in spring and 36% higher for plants at 185 µmol m-2 s-1 in spring compared to no ICL in spring. This increased autumn yield was caused by more fruiting laterals (less necrotic buds). It is concluded that management practices in spring can have a carryover effect on the autumn production. This is the first scientific paper on the potential for applying LED ICL in blackberries. Further research should focus on optimal intensity of ICL, positioning of supplementary lighting and economic feasibility.

Green light reduces elongation when partially replacing sole blue light independently from cryptochrome 1a.

Although green light is sometimes neglected, it can have several effects on plant growth and development. Green light is probably sensed by cryptochromes (crys), one of the blue light photoreceptor families. The aim of this study is to investigate the possible interaction between green and blue light and the involvement of crys in the green light response of plant photomorphogenesis. We hypothesize that green light effects on morphology only occur when crys are activated by the presence of blue light. Wild-type Moneymaker (MM), cry1a mutant (cry1a), and two CRY2 overexpressing transgenic lines (CRY2-OX3 and CRY2-OX8) of tomato (Solanum lycopersicum) were grown in a climate chamber without or with green light (30 µmol m-2  s-1 ) on backgrounds of sole red, sole blue and red/blue mixture, with all treatments having the same photosynthetic photon flux density of 150 µmol m-2  s-1 . Green light showed no significant effects on biomass accumulation, nor on leaf characteristics such as leaf area, specific leaf area, and chlorophyll content. However, in all genotypes, green light significantly decreased stem length on a sole blue background, whereas green light hardly affected stem length on sole red and red/blue mixture background. MM, cry1a, and CRY2-OX3/8 plants all exhibited similar responses of stem elongation to green light, indicating that cry1a, and probably cry2, is not involved in this green light effect. We conclude that partially replacing blue light by green light reduces elongation and that this is independent of cry1a.

LED and HPS Supplementary Light Differentially Affect Gas Exchange in Tomato Leaves.

Using light emitting diodes (LED) instead of conventionally used high pressure sodium (HPS) lamps as a supplemental light source in greenhouses results in a higher efficacy (µmol light per J electricity) and makes it possible to customize the light spectrum. To explore the effects of LED and HPS on gas exchange, thermal relations, photosynthesis, and water status of young tomato plants, seven genotypes were grown in a greenhouse under LED (95% red, 5% blue) or HPS lamps in four experiments differing in the fraction of lamp light over natural light. HPS lights emit a broader spectrum of red (40%), green-yellow (50%), blue (5%), and far-red (5%) and a substantial amount of infrared radiation (heat). Young tomato plants grown under LED showed lower leaf temperature and higher stomatal density, stomatal conductance (gs) and transpiration rate (E) than plants grown under HPS; this may be due to the different supplemental light spectrum. The young plants grown under LED tended to have increased photosynthetic capacity. Furthermore, the water stress indices CWSI and IG, which were obtained using thermal imaging, were positively correlated with gas exchange-derived gs and E, putting forward the use of thermal imaging for the phenotyping of transpiration. Under LED light, photosynthetic gas exchange was generally increased, which agreed with the water stress indices. The extent of this increase was genotype-dependent. All differences between LED and HPS were smaller in the experiments where the fraction of lamp light over natural light was smaller.

High Stomatal Conductance in the Tomato Flacca Mutant Allows for Faster Photosynthetic Induction.

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.

Far-red radiation stimulates dry mass partitioning to fruits by increasing fruit sink strength in tomato.

Far-red (FR) light promotes fruit growth by increasing dry mass partitioning to fruits, but the mechanism behind this is unknown. We hypothesise that it is due to an increased fruit sink strength as FR radiation enhances sugar transportation and metabolism. Tomato plants were grown with or without 50-80 µmol m-2  s-1 of FR radiation added to a common background 150-170 µmol m-2  s-1 red + blue light-emitting diode lighting. Potential fruit growth, achieved by pruning each truss to one remaining fruit, was measured to quantify fruit sink strength. Model simulation was conducted to test whether the measured fruit sink strength quantitatively explained the FR effect on dry mass partitioning. Starch, sucrose, fructose and glucose content were measured. Expression levels of key genes involved in sugar transportation and metabolism were determined. FR radiation increased fruit sink strength by 38%, which, in model simulation, led to an increased dry mass partitioned to fruits that quantitatively agreed very well with measured partitioning. FR radiation increased fruit sugar concentration and upregulated the expression of genes associated with both sugar transportation and metabolism. This is the first study to demonstrate that FR radiation stimulates dry mass partitioning to fruits mainly by increasing fruit sink strength via simultaneous upregulation of sugar transportation and metabolism.

Vertical Farming: Moving from Genetic to Environmental Modification.

Vertical farming (VF) is a novel plant production system that allows local production of high-quality fruits and vegetables for rapidly growing cities. VF offers a myriad of opportunities to move from genetic to environmental modification and to produce crops of guaranteed quality and quantity independent of weather, soil conditions, or climate change.

Adding Far-Red to Red-Blue Light-Emitting Diode Light Promotes Yield of Lettuce at Different Planting Densities.

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.

Dissecting the Genotypic Variation of Growth Responses to Far-Red Radiation in Tomato.

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.

Floral Induction in the Short-Day Plant Chrysanthemum Under Blue and Red Extended Long-Days.

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.