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.

Paradise by the far-red light: Far-red and red:blue ratios independently affect yield, pigments, and carbohydrate production in lettuce, Lactuca sativa.

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. Increasing 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.

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.

Short-term salt stress reduces photosynthetic oscillations under triose phosphate utilization limitation in tomato.

Triose phosphate utilization (TPU) limitation is one of the three biochemical limitations of photosynthetic CO2 assimilation rate in C3 plants. Under TPU limitation, abrupt and large transitions in light intensity cause damped oscillations in photosynthesis. When plants are salt-stressed, photosynthesis is often down-regulated particularly under dynamic light intensity, but how salt stress affects TPU-related dynamic photosynthesis is still unknown. To elucidate this, tomato (Solanum lycopersicum) was grown with and without sodium chloride (NaCl, 100 mM) stress for 13 d. Under high CO2 partial pressure, rapid increases in light intensity caused profound photosynthetic oscillations. Salt stress reduced photosynthetic oscillations in leaves initially under both low- and high-light conditions and reduced the duration of oscillations by about 2 min. Besides, salt stress increased the threshold for CO2 partial pressure at which oscillations occurred. Salt stress increased TPU capacity without affecting Rubisco carboxylation and electron transport capacity, indicating the up-regulation of end-product synthesis capacity in photosynthesis. Thus salt stress may reduce photosynthetic oscillations by decreasing leaf internal CO2 partial pressure and/or increasing TPU capacity. Our results provide new insights into how salt stress modulates dynamic photosynthesis as controlled by CO2 availability and end-product synthesis.

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.

Rapid spatial assessment of leaf-absorbed irradiance.

Image-based high-throughput phenotyping promises the rapid determination of functional traits in large plant populations. However, interpretation of some traits - such as those related to photosynthesis or transpiration rates - is only meaningful if the irradiance absorbed by the measured leaves is known, which can differ greatly between different parts of the same plant and within canopies. No feasible method currently exists to rapidly measure absorbed irradiance in three-dimensional plants and canopies. We developed a method and protocols to derive absorbed irradiance at any visible part of a canopy with a thermal camera, by fitting a leaf energy balance model to transient changes in leaf temperature. Leaves were exposed to short light pulses (30 s) that were not long enough to trigger stomatal opening but strong enough to induce transient changes in leaf temperature that was proportional to the absorbed irradiance. The method was successfully validated against point measurements of absorbed irradiance in plant species with relatively simple architecture (sweet pepper, cucumber, tomato, and lettuce). Once calibrated, the model was used to produce absorbed irradiance maps from thermograms. Our method opens new avenues for the interpretation of plant responses derived from imaging techniques and can be adapted to existing high-throughput phenotyping platforms.

Petunia as a model for MYB transcription factor action under salt stress.

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.

Corrigendum: Variation of photosynthetic induction in major horticultural crops is mostly driven by differences in stomatal traits.

[This corrects the article DOI: 10.3389/fpls.2022.860229.].

Vertical farming.

Ji et al. introduce vertical farming, a production-scale crop-growth system within an enclosed structure that maximizes space-use efficiency by growing plants vertically and precisely controls the plant environment.

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.

The association between the susceptibility to Botrytis cinerea and the levels of volatile and non-volatile metabolites in red ripe strawberry genotypes.

The relations between physical and chemical characteristics (e.g., color, firmness, volatile and non-volatile metabolites) of red ripe strawberry fruit and the natural spoilage caused by Botrytis cinerea were investigated. The spoilage rates differed between genotypes, and this was highly correlated over two successive years. Among seventeen genotypes, a more intense red coloration of the fruit skin was associated with a lower spoilage rate (r = -0.63). Additionally, weakly negative correlations were found between the levels of anthocyanins, ascorbic acid, malic acid and spoilage rates. No clear correlations were found between spoilage rates and soluble sugars, most volatiles, firmness and dry weight percentage. High levels of two volatile compounds, ethyl butanoate (r = 0.55) and 1-hexanol (r = 0.61), were correlated to high spoilage rates. These characteristics may assist strawberry breeders in selecting for genotypes with reduced susceptibility to B. cinerea.

Variation of Photosynthetic Induction in Major Horticultural Crops Is Mostly Driven by Differences in Stomatal Traits.

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.

Lack of Blue Light Regulation of Antioxidants and Chilling Tolerance in Basil.

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 F v /F m 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.

NaCl affects photosynthetic and stomatal dynamics by osmotic effects and reduces photosynthetic capacity by ionic effects in tomato.

NaCl stress affects stomatal behavior and photosynthesis by a combination of osmotic and ionic components, but it is unknown how these components affect stomatal and photosynthetic dynamics. Tomato (Solanum lycopersicum) plants were grown in a reference nutrient solution [control; electrical conductivity (EC)=2.3 dS m-1], a solution containing additional macronutrients (osmotic effect; EC=12.6 dS m-1), or a solution with additional 100 mM NaCl (osmotic and ionic effects; EC=12.8 dS m-1). Steady-state and dynamic photosynthesis, and leaf biochemistry, were characterized throughout leaf development. The osmotic effect decreased steady-state stomatal conductance while speeding up stomatal responses to light intensity shifts. After 19 d of treatment, photosynthetic induction was reduced by the osmotic effect, which was attributable to lower initial stomatal conductance due to faster stomatal closing under low light. Ionic effects of NaCl were barely observed in dynamic stomatal and photosynthetic behavior, but led to a reduction in leaf photosynthetic capacity, CO2 carboxylation rate, and stomatal conductance in old leaves after 26 d of treatment. With increasing leaf age, rates of light-triggered stomatal movement and photosynthetic induction decreased across treatments. We conclude that NaCl impacts dynamic stomatal and photosynthetic kinetics by osmotic effects and reduces photosynthetic capacity by ionic effects.

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.